contents pagesynergy.st-andrews.ac.uk/evolutionevolving/files/2019/03/... · 2019-03-28 ·...

Contents Page

Conference Welcome ......................................................................................................1

Important Information.....................................................................................................2

Invited Speakers...............................................................................................................5

Program At A Glance........................................................................................................8

Detailed Program...........................................................................................................10

Monday 1st April...................................................................................................10

Tuesday 2nd April..................................................................................................11

Wednesday 3rd April.............................................................................................14

Thursday 4th April.................................................................................................17

Poster Presentations......................................................................................................20

Delegate List ..................................................................................................................24

Abstracts: Invited Speakers............................................................................................31

Abstracts: Selected Talks................................................................................................37

Abstracts: Poster Presentations.....................................................................................99

1

Conference Welcome

Welcome to Churchill College in the beautiful city of Cambridge, home to Evolution

Evolving. The title of this meeting endeavours to highlight both that the evolutionary

process itself evolves over time and that evolutionary biology is a vibrant field of

enquiry with a theoretical framework that also evolves. Evolvability is just one of several

emerging themes in the relationship between development and evolution that

constitute the unique focus of this conference. Additional topics include the

evolutionary causes and consequences of developmental bias, plasticity, niche

construction and extra-genetic inheritance. Also distinctive of Evolution Evolving is the

inclusion of contributions to the history and philosophy of evolutionary biology, which

we believe are underappreciated but critically important to our field. We have tried

hard to ensure that the meeting features a balanced program of talks and poster

sessions, empirical and theoretical work, and a good mix of presentations from

researchers of all genders, career stages, and backgrounds. We hope that you enjoy

three wonderful days listening to marvelous talks and discussing evolutionary science,

and leave the meeting feeling both excited and inspired.

The organisers

Paul Brakefield Kevin Laland Tobias Uller

University of Cambridge University of St Andrews Lund University

Katrina Falkenberg Andrew Buskell

University of St Andrews University of Cambridge

2

Important Information

The Venue

A map of Churchill College can be found on page four. Larger paper maps of Churchill

College and Cambridge itself are available at the registration desk.

Main Concourse (C)

- registration

- welcome reception

- poster sessions

- morning and afternoon coffee breaks

Wolfson Hall (WH)

- morning and afternoon talk sessions

Fellow’s Dining Room (FDR)

- afternoon talk sessions

Churchill College Dining Hall (DH)

- lunches

- conference dinner

Seminar Room 6

- a quiet breakout space; this room is located close to staircase 37 in East Court

(denoted ‘E’ on the map) and directly opposite the Jock Colville Hall

Meals and Refreshments

Included in your registration fees are the welcome reception on Monday evening, as

well as morning and afternoon teas and fork buffet lunches on Tuesday, Wednesday

and Thursday. Limited drinks will be served at the poster sessions. Additional drinks can

be purchased from The Buttery (College bar), which is adjacent to the main concourse.

The Buttery serves light snacks and refreshments throughout the day.

Wifi

Eduroam is available on site. Alternatively, you can collect a wifi ticket from the

registration desk.

3

Social Media

We encourage the use of social media throughout the conference as

it is a powerful tool to share findings and engage with the scientific

community at large. Presenters: if you do not wish for your talk or a

particular slide to be tweeted, please indicate this with an image

such as the one to the right. Social media users: please respect the

wishes of presenters, and do not tweet images of posters without the

presenter’s permission.

#evoevolving is the conference hashtag and you can tag us with @evoevolving.

We recommend presenters and social media users read the social media guidelines of

the World Marine Mammal Conference 2019:

http://smmconference.org/socialmediaguidelines

Sponsors

We are very grateful for financial support from the John Templeton Foundation and The

Galton Institute.

https://twitter.com/search?f=tweets&vertical=default&q=%23evoevolving&src=typd&lang=en-gb

https://twitter.com/EvoEvolving?lang=en-gb

http://smmconference.org/socialmediaguidelines

5

Invited Speakers

Alexander Badyaev

Ecology and Evolutionary Biology, The University of Arizona, USA

Badyaev is a Professor of Evolutionary Biology. He addresses

conceptual questions at the intersection of evolution, ecology, and

development using a wide array of empirical and analytical

approaches in field and laboratory study systems.

Renee Duckworth


The ultimate goal of Duckworth’s work is to understand the link

between micro and macroevolutionary processes with specific focus

on the role of behavior in ecological and evolutionary feedbacks. To

achieve these goals, she integrates approaches from evolutionary

and physiological ecology using both large scale field experiments

and laboratory systems.

Laurel Fogarty

Max Planck Institute for Evolutionary Anthropology, Germany

Fogarty works on theory related to non-genetic inheritance. Most of

her work has a focus on cultural inheritance, cultural evolution, and

cultural niche construction. She also has a general interest in

theoretical niche construction.

Jukka Jernvall

Institute of Biotechnology, University of Helsinki, Finland

Jernvall’s research focuses on the developmental and ecological

drivers of phenotypic evolution. By integrating different scales and

methodologies, his group has used the mammalian dentition to

explore how morphology can inform us about evolution.

6

Alan C Love

Department of Philosophy and Minnesota Center for Philosophy of

Science, University of Minnesota - Twin Cities, USA

Love’s research focuses on conceptual issues in biology and has

concentrated on evolutionary developmental biology,

developmental biology, molecular biology, and paleontology. He

uses a combination of approaches to investigate a variety of

philosophical questions, including conceptual change, explanatory

pluralism, knowledge structure, reductionism, the nature of

historical science, and interdisciplinary epistemology.

Joanna Masel


Masel is a theoretical population geneticist and molecular

evolutionist. She studies a range of foundational questions in

evolution, including robustness and evolvability, the de novo birth

of proteins and their subsequent long-term change, and the

interplay between relative and absolute forms of competition.

Armin P Moczek

Department of Biology, Indiana University, USA

Moczek’s interests revolve broadly around the interplay of genetic,

developmental, and ecological mechanisms in the origin and

diversification of novel, complex traits. His work focuses on insects,

from beetles and their horns to fireflies and their bio-luminescent

lanterns.

Angela Potochnik

Department of Philosophy, University of Cincinnati, USA

Potochnik is an Associate Professor of Philosophy and the Director

of the Center for Public Engagement with Science at the University

of Cincinnati. Her research addresses the nature of science and its

successes, the relationships between science and the public, and

methods in population biology. She is the author of Idealizations and

the Aims of Science (2017) and coauthor of Recipes for Science: An

Introduction to Scientific Methods and Reasoning (2018).

7

Sean Rice

Department of Biological Sciences, Texas Tech University, USA

Rice’s research focuses on the mathematical principles underlying

evolutionary processes. He is particularly interested in the role of

stochasticity in the evolution of complex behavioural and

developmental systems.

Jessica Riskin

Department of History, Stanford University, USA

Riskin’s research interests centre upon the history of science,

politics and culture during the 17th to 19th centuries. She is currently

working on a set of connected research projects in the history of

evolutionary theory, including one on Lamarck and another on ideas

about human evolution.

8

Program At A Glance

Monday 1st April

18.00 - 19.45 Welcome reception (C)

Tuesday 2nd April

08.50 - 10.00 Conference welcome and plenary talk (WH)

10.00 - 10.30 Coffee Break (C)

10.30 - 12.10 Selected talks (WH)

12.10 - 13.30 Lunch (DH)

13.30 - 14.10 Keynote talks, two parallel sessions (WH, FDR)

14.10 - 15.10 Selected talks, two parallel sessions (WH, FDR)



17.20 - 19.00 Poster Session (C)

Wednesday 3rd April

09.00 - 10.00 Plenary talk (WH)



12.10 - 13.30 Lunch (DH)





17.20 - 19.00 Poster Session (C)

9

Thursday 4th April




12.10 - 13.30 Lunch (DH)





17.20 - 18.30 Free Time


19.45 – 23.00 Conference Dinner (DH)

WH = Wolfson Hall

FDR = Fellow’s Dining Room

C = Concourse

DH = Churchill College Dining Hall

10

Detailed Program

Monday 1st April

18.00 - 19.45 Registration

18.00 - 19.45 Welcome reception

11

Tuesday 2nd April

Wolfson Hall Morning

08.50 - 09.00 Conference Welcome: Kevin Laland

09.00 - 10.00

Plenary Talk: Renee Duckworth

Dynamic stability and robustness of ecological systems: a missing

link to evolutionary mechanisms?

The University of Arizona, USA

10.00 - 10.30 Coffee Break

10.30 - 12.10 General Session

10.30 - 10.50

Mycalesina in morphospace: how developmental bias shapes

evolutionary diversification in butterflies

Oskar Brattström, University of Cambridge, UK

10.50 - 11.10

Fighting about frequency

Karen Kovaka, Virginia Tech, USA

11.10 - 11.30

On leaders and followers: the evolution of phenotypic plasticity can

lead the evolution of genetic evolvability more easily than vice

versa

Miguel Brun-Usan, University of Southampton, UK

11.30 - 11.50

Bird nests evolve flexibly in response to egg characteristics

Catherine Sheard, University of St Andrews, UK

11.50 - 12.10

Adaptive evolution NOT driven by natural selection (?!?). On

individual responses, inclusive inheritance, and fleshing out the

second term of that famous Price equation

Pim Edelaar, University Pablo de Olavide, Spain

12.10 - 13.30 Lunch

12

Wolfson Hall Afternoon

13.30 - 14.10 Keynote Talk: Armin P Moczek

On the origins of novelty and diversity in development and

evolution: case studies on horned beetles

Indiana University, USA

14.10 - 17.20 Themed Session: Developmental Plasticity

14.10 - 14.30

DNA methylation lends plasticity to the coloration of an African

cichlid

Sebastian Alvarado, Queens College CUNY, USA

14.30 - 14.50

Unexplored maternal effect mediators, thyroid hormones, in an

eco-evolutionary context

Suvi Ruuskanen, University of Turku, Finland

14.50 - 15.10

Phenotypic plasticity facilitates colonization of savannas in

Mycalesina butterflies

Sridhar Halali, University of Cambridge, UK


15.40 - 16.00

Does thermal plasticity align with local adaptation? – An

interspecific comparison of wing shape and size in sepsid flies

Patrick Rohner, University of Zurich, Switzerland

16.00 - 16.20

Macroevolutionary dynamics of phenological plasticity

Stephen De Lisle, Lund University, Sweden

16.20 - 16.40

Developmental regulation and evolution of nutrition-responsive

growth: transcriptomic underpinnings of plasticity in horned

beetles

Sofia Casasa, Indiana University, USA

16.40 - 17.00

Involvement of maternal yolk thyroid hormones in the evolution of

life history traits in birds: a comparative analysis

Bin-Yan Hsu, University of Turku, Finland

17.00 - 17.20

Phenotypic divergence across spadefoot toad species mirrors

within-species consequences and mechanisms of developmental

plasticity

Christoph Liedtke, Doñana Biological Station, CSIC, Spain

17.20 - 19.00 Poster Session

13

Fellow’s Dining Room Afternoon

13.30 - 14.10 Keynote Talk: Angela Potochnik

Causal patterns and how our theories change

University of Cincinnati, USA

14.10 - 17.20 Themed Session: Philosophy of Biology

14.10 - 14.30

What the history of biology tells us about the Extended

Evolutionary Synthesis

Jan Baedke, Ruhr University Bochum, Germany

Daniel J Nicholson, Konrad Lorenz Institute for Evolution and

Cognition Research, Austria

14.30 - 14.50

Towards a novel unifying theory of the genotype concept

David Ricote Hernández, Centro Andaluz de Biología del Desarrollo,

CSIC-Universidad Pablo de Olavide-Junta de Andalucía, Spain

14.50 - 15.10

Rethinking cybernetics in contemporary theoretical biology

Flavia Fabris, Egenis, the Centre for the Study of Life Sciences,

University of Exeter, UK


15.40 - 16.00

The Extended Evolutionary Synthesis debate: some epistemological

and sociological dimensions

Alejandro Fábregas-Tejeda and Francisco Vergara-Silva, Instituto de

Biología, Universidad Nacional Autónoma de México, Mexico

16.00 - 16.20

Beyond standard views of reproduction: pregnant females as

individuals and their role in evolution

Laura Nuño de la Rosa and Arantza Etxeberria, Complutense

University of Madrid, Spain

16.20 - 16.40

Evolution of multicellularity: cheating done right

Walter Veit, University of Bristol, UK

16.40 - 17.00

The explanatory robustness of developmental types: a propensity

view

Cristina Villegas, Complutense University of Madrid, Spain

17.00 - 17.20

The cultural evolution of cultural evolution

Jonathan Birch, London School of Economics & Political Science, UK


14

Wednesday 3rd April


09.00 - 10.00

Plenary Talk: Alan C Love

Structuring knowledge in evolutionary biology

University of Minnesota - Twin Cities, USA



10.30 - 10.50

Evolution by natural induction

Richard Watson, University of Southampton, UK

10.50 - 11.10

Generalized pathways to specific outcomes: stress links maternal

environment and adaptive offspring trait variation

Ahva Potticary, The University of Arizona, USA

11.10 - 11.30

Evolution of migratory behavior by barnacle geese: the roles of

experience, memory and social learning

Thomas Oudman, NIOZ Netherlands Institute for Sea Research, The

Netherlands, and University of St Andrews, UK

11.30 - 11.50

Theory-based ecology and niche construction

Liz Pásztor, Biological Institute, Eötvös Loránd University, Hungary

11.50 - 12.10

The Hot Spring Hypothesis for the origin of life: implications for

niche construction theory in the first major transition in evolution

from non-life to Life

Bruce Damer, The Biota Institute, USA, and UC Santa Cruz, USA

12.10 - 13.30 Lunch

15


13.30 - 14.10 Keynote Talk: Jukka Jernvall

To walk or not to walk the line of a developmental bias

University of Helsinki, Finland

14.10 - 16.00 Themed Session: Developmental Bias

14.10 - 14.30

Developmental bias and morphological convergence in Anolis

lizards

Nathalie Feiner, Lund University, Sweden

14.30 - 14.50

Measuring developmental bias: quantitative genetics meets

developmental biology

Lisandro Milocco, University of Helsinki, Finland

14.50 - 15.10

The evolutionary potential of environmentally induced

phenotypes

Reinder Radersma, Lund University, Sweden


15.40 - 16.00

Developmental interactions established during speciation leave a

lasting trace in descendent populations

Laura Mears, University of Southampton, UK

16.00 - 17.20 Themed Session: Evolutionary Innovation

16.00 - 16.20

What kind of evolutionary novelty (if any) could be language?

Salvatore Ivan Amato and Giuliana Pulvirenti, University of

Messina, Italy

16.20 - 16.40

It’s a horn, a wing, a helmet: the role of wing serial homologs in

insect innovations

Yonggang Hu, Indiana University, USA

16.40 - 17.00

Beyond Darwin: how every individual adapts to its internal

perturbations?

Yoav Soen, Weizmann Institute of Science, Israel

17.00 - 17.20

How the beetle head got its horns: integrating a novel trait into a

conserved trait complex

David M Linz, Indiana University, USA


16


13.30 - 14.10 Keynote Talk: Laurel Fogarty

The inheritance and heritability of niche constructed traits


14.10 - 16.20 Themed Session: Niche Construction

14.10 - 14.30

Niche construction affects the consistency and strength of natural

selection

Andrew Clark, University of St Andrews, UK

14.30 - 14.50

Linking coral morphological traits and reef fish community structure

Luisa Fontoura, Macquarie University, Australia

14.50 - 15.10

Microbial community structure and function driven by infaunal niche

construction in intertidal sediments

Adam Wyness, University of St Andrews, UK


15.40 - 16.00

Keeping close neighbours, corals are spatially clustered within and

across species

Maria Dornelas, University of St Andrews, UK

16.00 - 16.20

Holed-up: nesting in a cavity as an evolutionary driver of parental

care in fishes and birds

Karina Vanadzina, University of St Andrews, UK

16.20 - 17.20 Themed Session: Cultural Evolution

16.20 - 16.40

Trait specialisation, innovation and the evolution of culture in

changing environments

Dominik Deffner, Max Planck Institute for Evolutionary Anthropology,

Germany

16.40 - 17.00

A New Caledonian crow's-eye-view on how to develop probing

intelligence: the evolution of tool-use and tool-making as embodied,

distributed and socially situated perception action processes

Joanna Fairlie, Independent Scholar

17.00 - 17.20

Generalized Darwinism revisited: how a new synthesis changes our

view on cultural evolution

Karim Baraghith, Heinrich-Heine University, Germany


17

Thursday 4th April


09.00 - 10.00

Plenary Talk: Alexander Badyaev

Control theory in evolution




10.30 - 10.50

Stress-induced mutagenesis systems and the adaptive immunity

Bartlomiej Swiatczak, University of Science and Technology of

China, China

10.50 - 11.10

Phenotypic integration as both developmental bias and

developmental facilitation of evolutionary diversification: empirical

examples from teeth, toes, and torsos

Kathryn Kavanagh, University of Massachusetts Dartmouth, USA

11.10 - 11.30

Can plasticity create irreversible constraints?

Alfredo Rago, University of Southampton, UK

11.30 - 11.50

Cultural evolution, revolution and transmission of humpback whale

song

Ellen C Garland, University of St Andrews, UK

11.50 - 12.10

Niche construction: resolving the process debate

Tim Lewens, University of Cambridge, UK

12.10 - 13.30 Lunch

18


13.30 - 14.10 Keynote Talk: Joanna Masel

Mutational and pre-mutational shaping of adaptation


14.10 - 17.20 Themed Session: Evolvability

14.10 - 14.30

Comprehensive transcriptomics tracks changes in gene expression

and coding sequences during C4 evolution

Lamiaa A Munshi, The University of Sheffield, UK

14.30 - 14.50

Computational complexity is an ultimate constraint on evolution

Artem Kaznatcheev, University of Oxford, UK

14.50 - 15.10

Population-level variation in sperm-mediated paternal effects in

threespined sticklebacks

Jennifer Hellmann, University of Illinois Urbana-Champaign, USA


15.40 - 16.00

The effect of parental care on population evolvability

Sonia Pascoal, University of Cambridge, UK

16.00 - 16.20

The thrifty watchmaker

Michael Lachmann, Santa Fe Institute, USA

16.20 - 16.40

Variable phenotypic plasticity and its consequences for

evolutionary potential

Franziska Brunner, University of Liverpool, UK

16.40 - 17.00

Comparative genomics of convergent evolution in the emergence

of complex traits

Claudio L Flores Martinez, University of Hamburg, Germany

17.00 - 17.20

Epigenetic potential in native and introduced populations of house

sparrows (Passer domesticus)

Haley Hanson, University of South Florida, USA

17.20 - 18.30 Free time

18.30 - 19.30 Plenary Talk: Jessica Riskin

The restless machinery of life

Stanford University, USA

19.45 - 23.00 Conference Dinner

19


13.30 - 14.10 Keynote Talk: Sean Rice

How to make general theories of evolution applicable to complex

special cases

Texas Tech University, USA

14.10 - 16.00 Themed Session: Mathematical Models

14.10 - 14.30

Why does natural selection favour the transgenerational

inheritance of social adversity?

Bram Kuijper, University of Exeter, UK

14.30 - 14.50

Trait mean-variance relationships impact speed of adaptation

Rebecca Hoyle, University of Southampton, UK

14.50 - 15.10

The influence of multi-level demographic and ecological feedbacks

in the evolution of cooperation

Antonio MM Rodrigues, University of Cambridge, UK


15.40 - 16.00

The evolution of teaching and anti-teaching

James Allen, University of Cambridge, UK

16.00 - 17.20 Themed Session: Inclusive Inheritance

16.00 - 16.20

Evolution of small RNA production under fluctuating

environmental conditions

Willian TAF Silva, Uppsala University, Sweden

16.20 - 16.40

Inheritance of acquired traits - implications for evolution?

Sophie Juliane Veigl, University of Vienna, Austria

16.40 - 17.00

Egg development plasticity and the transmission of maternal

effects in Daphnia

Stewart Plaistow, University of Liverpool, UK

17.00 - 17.20

Environmental heterogeneity mediates the evolution of trans-

generational effects on fitness

Martin Lind, Uppsala University, Sweden

20

Poster Presentations

P01 Expanding the conceptual scope of niche construction theory

Bendik Hellem Aaby, KU Leuven, Belgium

P02 Could a major evolutionary transition have happened in human evolution? Paulo C Abrantes, University of Brasília, Brazil

P03 Dialectics of evolution; epistemology versus practice

Ozan A Altinok, WWU Münster, Germany

P04 Overprinting and the evolvability of novel protein-coding genes

Zachary Ardern, Technical University of Munich, Germany

P05 Morphological divergence in Drosophila ananassae due to nutritional

variations through changes in allometric relationship and trait size

Bhumika, Banaras Hindu University, India

P06 Coral-built structural complexity enhances coral recruitment: niche

construction in action?

Viviana Brambilla, University of St Andrews, UK

P07 When are more experiments necessary? Using evolutionary considerations in

investigating animal minds

Simon Brown, Columbia University, USA

P08 The evolution of unicellularity in a multicellular world

Charlie Cornwallis, Lund University, Sweden

P09 Enactive evolution

Amanda Corris, University of Cincinnati, USA

P10 Behemoth: Henri Bergson's and Gilles Deleuze's vitalism, a proto evolutionary

biology

Jack Robert Edmunds-Coopey, University of Durham, UK

P11 Effects of phenotypic robustness on adaption and evolvability

Giuseppe Fusco, University of Padova, Italy

P12 Spadefoot toad tadpoles incur maintenance costs of developmental plasticity

in the form of increased oxidative stress

Ivan Gomez-Mestre, Doñana Biological Station, CSIC, Spain

P13 The evolution of barriers to exploitative behaviors

Jonathan R Goodman, University of Cambridge, UK

21

P14 Three lessons of Bergson's creative evolution in the wake of transhuman

evolution

Daina Habdankaite, Vilnius University, Lithuania, and Turin University, Italy

P15 Evolution of phenotypic complexity: a developmental perspective

Pascal Felix Hagolani, University of Helsinki, Finland

P16 Timing of maternal exposure to toxic cyanobacteria and offspring fitness in

Daphnia magna: implications for the evolution of anticipatory maternal effects

Alexander Hegg, Lund University, Sweden

P17 Studying the evolutionary process with genetic algorithms

Edith Invernizzi, University of St Andrews, UK

P18 The evolutionary origin of our moral beliefs

Man-Him Ip, University of Birmingham, UK

P19 Genetic assimilation in the fossil record: phenotypic plasticity and

accommodation in Cambrian arthropods

Illiam Jackson, Lund University, Sweden

P20 Extended cognitive systems, meaning, and evolutionary transitions

Hilton F Japyassú, Universidade Federal da Bahia, Brazil

P21 Mechanisms of hormone mediated parent-offspring conflict

Neeraj Kumar, University of Groningen, The Netherlands, and Max Planck

Institute for Ornithology, Germany

P22 The impact of individual immune experience on niche construction in red flour

beetles, Tribolium castaneum

Lai Ka Lo, University of Münster, Germany

P23 Trait-based theory leads to generality across successional systems

Jessica Haghkerdar, University of St Andrews, UK

P24 From anisogamy to intrasexual competition

Ivain Martinossi, Uppsala University, Sweden

P25 Diet-based developmental plasticity in a freshwater isopod

Blake Matthews, Eawag, Switzerland

P26 Global human niche construction: an evolutionary trap?

Andra Meneganzin, Università degli Studi di Padova, Italy

P27 A weapons-testes tradeoff in males extends to female traits

Christine Miller, University of Florida, USA

22

P28 Assessing adaptive and non-adaptive evolution of the glycolytic enzyme

triosephosphate isomerase

Ricardo Muñiz Trejo, National Autonomous University of Mexico, Mexico

P29 The evolution of hierarchical structure: separating the causes and

consequences of evolvability

Frederick J Nash, University of Southampton, UK

P30 Spandrels and trait delimitation: no such thing as “architectural constraint”

Mark E Olson, Universidad Nacional Autónoma de México, Mexico

P31 With a little help from my friends: the role of the microbiota in dung beetle

diversification

Erik Parker, Indiana University, USA

P32 Transposable elements: the unexpected junk that shaped genomes and

challenged evolutionary biology

Valentina Peona, Uppsala University, Sweden

P33 Weismann, Crick, and others: flows of information in evolution

Arnaud Pocheville, CNRS and Université Paul Sabatier, France

P34 Delineating the boundaries of adaptation by tracing histories of environmental

induction

Ahva Potticary, The University of Arizona, USA

P35 The interaction of learning and evolution can enable adaptive behaviours that

are not presupposed in the learning ability

David Prosser, University of Southampton, UK

P36 Naturalising the language faculty: the role of structures and developmental

processes

Giuliana Pulvirenti, University of Messina, Italy

P37 Teleology and evolution

Rasmus Sandnes Haukedal, Durham University, UK

P38 The evolution of adaptive epigenetic inheritance

Zuzana Sekajova, Uppsala University, Sweden

P39 Unique evolutionary properties of plastic traits: an empirical study of erosion

of phenotypic plasticity under relaxed selection

Helen Spence-Jones, University of St Andrews, UK

P40 A culturally driven plasticity hypothesis for the evolution of human life course

Francesco Suman, Independent Scholar

23

P41 Unique, essential, and unknown: the enigma of taxonomically restricted,

essential genes of unknown function

Change Tan, University of Missouri, USA

P42 The evolution of relatedness and evolutionary transitions in individuality

Christoph Thies, University of Southampton, UK

P43 Deconstructing reef complexity: relating coral diversity, rugosity and fractal

dimension

Damaris Torres-Pulliza, Macquarie University, Australia

P44 Cultural evolution and neuroscience: a pathway to integration

Ryutaro Uchiyama, London School of Economics, UK

P45 Lost (and found) in translation: circulation of EES-related concepts in two

scientific communities in Mexico

Francisco Vergara-Silva and Alejandro Fábregas-Tejeda, Universidad Nacional

Autónoma de México, Mexico

P46 Testing the flexible stem hypothesis in stickleback fish: is ancestral

morphological plasticity adaptive?

Matthew Wund, The College of New Jersey, USA

P47 Quantifying niche constructing traits in reef corals

Kyle Zawada, University of St Andrews, UK

P48 Two senses of biological possibility

Jason Zinser, University of Wisconsin - Stevens Point, USA

P49 Plasticity in mouthpart length in response to developmental diet in leaf-footed

bugs

Sara Zlotnik, University of Florida, USA

P50 Evolution and development of polyphenisms: a mechanistic model based on

bistable switches

Roman Zug, Lund University, Sweden

24

Delegate List Aaby, Bendik Hellem KU Leuven P01

Abrantes, Paulo C University of Brasília P02

Allen, James University of Cambridge FDR, 4th

Altinok, Ozan A WWU Münster P03

Alvarado, Sebastian Queens College CUNY WH, 2nd

Amato, Salvatore Ivan University of Messina WH, 3rd

Ardern, Zachary Technical University of Munich P04

Armfield, Roxanne University of Cambridge

Arnold, Kevin John Templeton Foundation

Badyaev, Alex The University of Arizona WH, 4th

Baedke, Jan Ruhr University Bochum FDR, 2nd

Banzhaf, Wolfgang Michigan State University

Baraghith, Karim Heinrich Heine University Düsseldorf FDR, 3rd

Beaudoin, Chris University of Cambridge

Berkemeier, Francisco University College London

Bhumika Banaras Hindu University P05

Birch, Jonathan London School of Economics FDR, 2nd

Bocedi, Greta University of Aberdeen

Brakefield, Paul University of Cambridge

Brambilla, Viviana University of St Andrews P06

Brattström, Oskar University of Cambridge WH, 2nd

Brown, Simon Columbia University P07

Brun-Usan, Miguel University of Southampton WH, 2nd

Brunner, Franziska University of Liverpool WH, 4th

Buskell, Andrew University of Cambridge

Campo Bes, Israel University of Valencia

Cano, Hugo University College London

Casasa, Sofia Indiana University WH, 2nd

Catherall, Andrew University of Cambridge

Chauvet, Adrien The University of Sheffield

25

Clark, Andrew University of St Andrews FDR, 3rd

Collins-Attwood, Mairenn University of Cambridge

Cornwallis, Charlie Lund University P08

Corris, Amanda University of Cincinnati P09

Damer, Bruce The Biota Institute, UC Santa Cruz WH, 3rd

De Lisle, Stephen Lund University WH, 2nd

De Sanctis, Bianca University of Cambridge

Deffner, Dominik Max Planck Institute for Evolutionary

Anthropology

FDR, 3rd

Dobreva, Mariya Imperial College London

Dornelas, Maria University of St Andrews FDR, 3rd

Duckworth, Renee The University of Arizona WH, 2nd

Edelaar, Pim University Pablo de Olavide WH, 2nd

Edmunds-Coopey, Jack Robert University of Durham P10

Etxeberria, Arantza University of the Basque Country FDR, 2nd

Fábregas-Tejeda, Alejandro Universidad Nacional Autónoma de

México

FDR, 2nd

Fabris, Flavia University of Exeter FDR, 2nd

Fairlie, Joanna Independent Scholar FDR, 3rd

Falkenberg, Katrina J University of St Andrews

Falzone, Alessandra University of Messina

Faria, Vitor G University of Basel

Feiner, Nathalie Lund University WH, 3rd

Feldman, Marcus Stanford University

Fjågesun, Tonje University of Cambridge

Flores Martinez, Claudio L University of Hamburg WH, 4th

Fogarty, Laurel Max Planck Institute for Evolutionary

Anthropology

FDR, 3rd

Fontoura, Luisa Macquarie University FDR, 3rd

Fusco, Giuseppe University of Padova P11

Gallardo Pinedo, Adrian University of Valencia

Garland, Ellen C University of St Andrews WH, 4th

George, Alison New Scientist

26

Gomez-Mestre, Ivan Doñana Biological Station, CSIC P12

Goodman, Jonathan R University of Cambridge P13

Goymer, Patrick Nature

Groothuis, Ton University of Groningen

Habdankaite, Daina Vilnius University, Turin University P14

Haghkerdar, Jessica University of St Andrews P23

Hagolani, Pascal Felix University of Helsinki P15

Halali, Sridhar University of Cambridge WH, 2nd

Halina, Marta University of Cambridge

Halle, Stefan Friedrich Schiller University Jena

Hanson, Haley University of South Florida WH, 4th

Hegg, Alexander Lund University P16

Helanterä, Heikki University of Oulu

Hellmann, Jennifer University of Illinois Urbana-

Champaign

WH, 4th

Henderson, Tara University of Cambridge

Hoyle, Rebecca University of Southampton FDR, 4th

Hsu, Bin-Yan University of Turku WH, 2nd

Hu, Yonggang Indiana University WH, 3rd

Invernizzi, Edith University of St Andrews P17

Ip, Man-Him University of Birmingham P18

Jackson, Illiam Lund University P19

Japyassú, Hilton F Universidade Federal da Bahia P20

Jernvall, Jukka University of Helsinki WH, 3rd

Johnstone, Rufus University of Cambridge

Kavanagh, Kathryn University of Massachusetts

Dartmouth

WH, 4th

Kaznatcheev, Artem University of Oxford WH, 4th

Kilner, Rebecca University of Cambridge

Kovaka, Karen Virginia Tech WH, 2nd

Kozielska, Magdalena University of Groningen

Kuijper, Bram University of Exeter - Penryn Campus FDR, 4th

27

Kumar, Neeraj University of Groningen,

Max Planck Institute for Ornithology

P21

Lachmann, Michael Santa Fe Institute WH, 4th

Laland, Kevin N University of St Andrews

Lazarus, John Newcastle University

Leitao, Alex University of Cambridge

Lewens, Tim University of Cambridge WH, 4th

Liedtke, Christoph Doñana Biological Station, CSIC WH, 2nd

Lind, Martin Uppsala University FDR, 4th

Linz, David M Indiana University WH, 3rd

Livnat, Adi University of Haifa

Lo, Lai Ka University of Münster P22

Love, Alan C University of Minnesota - Twin Cities WH, 3rd

Madin, Joshua S University of Hawaii

Maeso, Ignacio Andalusian Centre for Developmental

Biology

Mahen, Beeshman University of Cambridge

Marconi, Aleksandra University of Cambridge

Martin, Marty University of South Florida

Martinez, Pedro Universitat de Barcelona

Martinossi, Ivain Uppsala University P24

Masel, Joanna The University of Arizona WH, 4th

Matthews, Blake Eawag P25

Mears, Laura University of Southampton WH, 3rd

Méndez Salinas, Emiliano University of Groningen

Meneganzin, Andra Università degli Studi di Padova P26

Merleau-Ponty, Noémie University of Cambridge

Miller, Christine University of Florida P27

Milocco, Lisandro University of Helsinki WH, 3rd

Moczek, Armin P Indiana University WH, 2nd

Muñiz Trejo, Ricardo National Autonomous University of

Mexico

P28

Munshi, Lamiaa A The University of Sheffield WH, 4th

28

Nash, Frederick J University of Southampton P29

Nelson, Paul Biola University

Nicholson, Daniel J Konrad Lorenz Institute for Evolution

and Cognition Research

FDR, 2nd

Nuño de la Rosa, Laura Complutense University of Madrid FDR, 2nd

Odling-Smee, John Mansfield College - Oxford

Odling-Smee, Ros University of Oxford - Retiree

Olson, Mark E Universidad Nacional Autónoma de

México

P30

Oudman, Thomas NIOZ Netherlands Institute for Sea

Research, University of St Andrews

WH, 3rd

Padilla Iglesias, Cecilia University of Cambridge

Page, Karen M University College London

Pagnotta, Murillo University of St Andrews

Parker, Erik Indiana University P31

Pascoal, Sonia University of Cambridge WH, 4th

Pásztor, Liz Eötvös University WH, 3rd

Paternò, Orazioluca Università degli Studi di Torino

Paterson, David University of St Andrews

Peona, Valentina Uppsala University P32

Perera, Roly Alan Turing Institute

Picao-Osorio, Joao Institute of Biology of the École

Normale Supérieure

Plaistow, Stewart University of Liverpool FDR, 4th

Pocheville, Arnaud CNRS, Université Paul Sabatier P33

Potochnik, Angela University of Cincinnati FDR, 2nd

Potticary, Ahva The University of Arizona WH, 3rd, P34

Pretelli, Ilaria Max Planck Institute for Evolutionary

Anthropology

Prosser, David University of Southampton P35

Pulvirenti, Giuliana University of Messina WH, 3rd, P36

Radersma, Reinder Lund University WH, 3rd

Rago, Alfredo University of Southampton WH, 4th

29

Ratikainen, Irja Norwegian University of Science and

Technology

Renton, Jessie University College London

Retzlaff, Nancy MPI for Mathematics in the Sciences,

Leipzig University

Rice, Sean Texas Tech University FDR, 4th

Ricote Hernández, David Andalusian Centre for Developmental

Biology

FDR, 2nd

Riederer, Jana University of Groningen

Riskin, Jessica Stanford University WH, 4th

Robin, Nicolas University of Teacher Education

St Gallen

Rodrigues, Antonio MM University of Cambridge FDR, 4th

Rohner, Patrick University of Zurich WH, 2nd

Ruuskanen, Suvi University of Turku WH, 2nd

Sandnes Haukedal, Rasmus Durham University P37

Santos, Emília University of Cambridge

Scherrer, Raphaël University of Groningen

Sekajova, Zuzana Uppsala University P38

Sheard, Catherine University of St Andrews WH, 2nd

Silva, Willian TAF Uppsala University FDR, 4th

Soen, Yoav Weizmann Institute of Science WH, 3rd

Spence-Jones, Helen University of St Andrews P39

Stocks, Miranda IST Austria

Suman, Francesco Independent Scholar P40

Swiatczak, Bartlomiej University of Science and Technology

of China

WH, 4th

Tan, Change University of Missouri P41

Thies, Christoph University of Southampton P42

Torres-Pulliza, Damaris Macquarie University P43

Toyokawa, Wataru University of Konstanz

Tschol, Maximilian University of Aberdeen

Tucker, Roger Sonocent Ltd

30

Uchiyama, Ryutaro London School of Economics P44

Uller, Tobias Lund University

van den Engel, Peter Independent Scholar

van Eldijk, Timo University of Gronignen

Vanadzina, Karina University of St Andrews FDR, 3rd

Vasilyeva, Zinaida Technical University of Munich

Veigl, Sophie Juliane University of Vienna FDR, 4th

Veit, Walter University of Bristol FDR, 2nd

Vergara-Silva, Francisco Universidad Nacional Autónoma de

México

FDR, 2nd, P45

Villegas, Cristina Complutense University of Madrid FDR, 2nd

Vinterstare, Jerker Lund University

Vykuntam, Vinit University of Cambridge

Wason, Paul John Templeton Foundation

Watson, Richard A University of Southampton WH, 3rd

While, Geoff University of Tasmania

White, Roger Memorial University of Newfoundland

Wilkins, Adam S Humboldt University Berlin

Winder, Isabelle Bangor University

Wund, Matthew The College of New Jersey P46

Wyness, Adam University of St Andrews FDR, 3rd

Yen, Eugenie University of Cambridge

Zawada, Kyle University of St Andrews P47

Zinser, Jason University of Wisconsin - Stevens Point P48

Zlotnik, Sara University of Florida P49

Zug, Roman Lund University P50

WH = talk in Wolfson Hall, date

FDR = talk in Fellow’s Dining Room, date

P = Poster

31

Abstracts: Invited Speakers

Dynamic stability and robustness of ecological systems: a missing link to evolutionary

mechanisms?

Renee Duckworth


Ecosystems are often viewed as self-organizing systems that remain stable in their

outputs. Their pattern of dynamic stability is remarkably similar to patterns at the

macroevolutionary scale in species’ stasis. Yet, it is unclear whether the types of

dynamics that produce stability in ecosystems are causally linked to patterns of species’

stasis, and, if so, what are the mechanisms underlying both? I suggest that ecological

communities are in a state of dynamic stability which is maintained by the active

responses of organisms to environmental change through processes such as niche

construction, adaptive plasticity and habitat selection. As such, persistent evolutionary

change may only occur when there is a major disruption of the ecological community

as a whole. To show how such dynamics might be studied and how a body of work can

be reinterpreted with this novel framework, I draw on my work on the population and

community dynamics of cavity nesting birds. I use this example to illustrate how a

deeper understanding of the mechanisms underlying dynamic stability may be crucial

to predicting the two most common outcomes of environmental change: whether

populations will go extinct or evolve.

32

On the origins of novelty and diversity in development and evolution: case studies on

horned beetles

Armin Moczek


Understanding how development and ecology shape organismal evolution is a central

goal of evolutionary developmental biology. In my presentation I highlight a class of

traits and organisms that are emerging as new models in evo devo and eco devo

research: beetle horns and horned beetles. Horned beetles are morphologically diverse,

ecologically rich, and developmentally and genetically increasingly accessible. Recent

studies have begun to take advantage of these attributes and are starting to link the

microevolution of horned beetle development to the macroevolution of novel, complex

traits, and to identify the genetic, developmental, and ecological mechanisms, and the

interactions between them, that mediate organismal innovation and diversification in

natural populations. Here, I review the most significant recent findings and their

contributions to current frontiers in evolutionary developmental biology. In particular,

I explore the role of developmental mechanisms in biasing and facilitating innovation in

evolution and the significance of host-symbiont interactions and niche construction in

ecological radiations.

Causal patterns and how our theories change

Angela Potochnik


In evolutionary biology, attention to causes—or mechanism—is common, as is debate

about which causes are important. In this talk, I motivate the idea that scientists aim to

identify not just causes, but what I call causal patterns. Focus on a given causal pattern

is motivated by a combination of causal facts and the nature of the research priorities.

This framing changes how we should think about theory development, including the

development of evolutionary theory. Theory change can be motivated in two different

ways: by the discovery that a causal pattern fails to obtain, or by a shift in research

priorities toward a focus on different causal patterns. On the other hand, this view also

predicts the maintenance of multiple accounts of the same phenomena, accounts that

may seem incompatible but that capture different causal patterns, of interest to

different research programs. I develop these points with attention to the relationship

between developmental biology and evolutionary theory.

33

Structuring knowledge in evolutionary biology

Alan C Love

University of Minnesota - Twin Cities, USA

Ongoing debates about the possibility and status of an extended evolutionary synthesis

often involve disagreements about the nature and significance of particular phenomena

(e.g., developmental plasticity) or mechanisms (e.g., phenotypic accommodation).

However, assumptions regarding how to structure knowledge about phenomena and

mechanisms in evolutionary biology remain largely implicit in these conversations. I

argue that greater attention to these assumptions—and the expectations they foster—

has the potential to reveal the conceptual basis of some existing controversies about

theoretical frameworks in evolutionary biology and provide novel ways of structuring

our evolving knowledge for future interdisciplinary inquiry.

To walk or not to walk the line of a developmental bias

Jukka Jernvall


Developmental processes can influence, or even bias phenotypic variation available for

natural selection. While developmental biases are perhaps generally acknowledged,

explanations of the specific impacts of development on evolution are easily little more

than ‘just so stories’. To provide more disciplined predictions about the effects of

development on evolution, developmental models or rules can be used. But what does

it mean when the data does not obey a developmental rule? I will discuss a

development-based ‘inhibitory cascade’ rule that predicts the evolution of mammalian

tooth proportions. Whereas many mammalian species appear to vary as predicted by

the inhibitory cascade rule, there are also exceptions. These exceptions can be argued

to disprove the inhibitory cascade, but the exceptions themselves should also be

examined critically. I will show how experimentally breaking the inhibitory cascade

uncovers the processes underlying the rules, and how evolutionary exceptions may not

be that exceptional after all.

34

The inheritance and heritability of niche constructed traits

Laurel Fogarty, Michael J Wade


A central tenet of niche construction theory is that organisms, through their own

actions, can alter the selective pressures to which they are subject and, in turn, change

the rates and perhaps trajectories of their species’ evolution. However, the ways in

which niche construction changes not just the selection on a trait but the structure of

the inheritance of that trait are poorly understood. Here we describe a quantitative

genetic model of niche construction that describes the inheritance patterns for a niche-

constructing trait and an associated niche-construction-sensitive trait. We show how

the response to selection can differ from that which would be expected in the absence

of niche construction as a result of changes to associated breeding values. We compare

these models to existing and closely related models of indirect genetic effects and

discuss the implications for our understanding of the heritability of niche constructing

traits and the importance of niche construction in evolutionary theory in general.

Control theory in evolution

Alexander Badyaev


With a few notable exceptions from neurobiology, cytology, and physiology, there is

little cross-talk between evolutionary theory and control theory of dynamic systems.

Yet, both fields are intensely focused on essentially an identical set of problems – on

reconciling stability and change, continuity and discreetness, innovability and

complexity, and inherency and contingency. Both fields aim to resolve these problems

by predicting the distribution and kinds of “controls” that either change or regulate

complex systems. As a result, we now have a set of parallel solutions to essentially the

same problems but from largely non-overlapping starting points, backgrounds, and

motivations. I will review these solutions, their empirical manifestations, and the

insights they provide into some of the most long-standing issues in evolutionary biology,

including the most enigmatic of them all – the relationship between form and function

in development and inheritance.

35

Mutational and pre-mutational shaping of adaptation

Joanna Masel


Evolvability depends on both the quantity and quality of heritable variants. For

polygenic traits, the quality of the mutational neighborhood is a more important

determinant of a population’s evolvability than the spread of the population across a

neutral genotype network. A first approximation of mutational quality is the ratio

between the two modes of the distribution of fitness effects, which tend either to

reduce fitness to zero, or have weak effects, but are rarely in between. If developmental

errors in the present mimic the effects of future mutations, selection can act pre-

mutationally to make the relative frequencies of these two modes more favorable. Data

on the cryptic sequences beyond stop codons suggests that highly expressed cryptic

sequences have experienced more such pre-mutational shaping. Evolutionary

capacitance to exploit such pre-screened sources of variation can either evolve through

capacitance widgets such as the yeast [PSI+] prion, or can emerge non-adaptively.

Capacitance makes the crossing of fitness valleys far more likely. Epimutations differ

from ordinary mutations in their rates – they are typically not just higher, but also lack

a strong excess of loss of function over gain of function – but are otherwise analyzable

by conventional theoretical population genetic approaches. The relative availability of

different kinds of beneficial mutation is known to shape adaptation when the product

of the beneficial mutation rate and census population size UN<1. When UN>1, clonal

interference is expected to make differences in the selection coefficient s much more

important than differences in U. However, the phenomenon of mutation-driven

adaptation re-emerges when U>s.

36

How to make general theories of evolution applicable to complex special cases

Sean Rice

Texas Tech University, USA

Much of evolutionary theory consists of special case models; built around simplifying

assumptions that make them tractable, but limit their applicability outside of the

particular systems for which they were built. We also have some very general results,

such as the Price equation, that are mathematically simple and in principle hold true for

all evolving systems. In practice, however, it is often difficult to apply these results to

particular systems of interest, since it is not clear how the details of complex biological

systems translate into the few terms of the general equation. I will show how new

mathematical tools allow us to build truly general theories -- requiring no simplifying

assumptions -- that are nonetheless applicable to the biological details of particular

systems. I will illustrate this approach by showing how it gives insight into the evolution

of complex cooperation, and helps us to understand large scale patterns in the

evolution of multicellularity.

The restless machinery of life

Jessica Riskin

Stanford University, USA

Ever since the beginnings of modern science in the 17th century, philosophers and

scientists have used clockwork and related machinery as a metaphor to describe the

universe and the natural world. The machinery model generally makes one think of

something rigid, rote, constrained, something that ticks and whirs along without

changing. However, hidden within this centuries-old clockwork-machinery metaphor is

a tradition of people who have understood artificial machinery, and nature, very

differently: as restless, responsive, and dynamic. These renegades include the German

philosopher Leibniz who first described living things as “organisms,” and who said living

organisms resembled clocks in their “restlessness”; the French naturalist Lamarck who

was the first to present a theory of what we now call evolution; and the Austrian

pioneer in quantum physics, Erwin Schrödinger, whose landmark 1944 manifesto What

is Life was foundational to the new field of molecular biology. All three compared living

beings to clockwork machinery but all three specified that they meant restless clocks.

The talk will explore the history of restless machinery as a model for living things.

37

Abstracts: Selected Talks

Mycalesina in morphospace: how developmental bias shape evolutionary

diversification in butterflies

Oskar Brattström, Kwaku Aduse-Poku, Erik van Bergen, Vernon French, Paul M

Brakefield

University of Cambridge, UK

The developmental mechanisms that generate morphology can in theory bias the

independent evolution of traits sharing ontogenetic pathways, making certain

evolutionary changes more likely than others. The eyespots commonly found on

butterfly wings each have concentric rings of differing colours and these classic

examples of serially repeated pattern elements have been a focus for evo-devo

research. Previous work on a model species of Mycalesina butterfly, Bicyclus anynana,

has provided insights into the developmental control of individual eyespots using

experimental evolution. These studies showed that the relative size of a eyespots on

the same wing surface is highly flexible, whereas they are resistant to diverging in

colour-composition, presumably due to shared developmental process. Here we survey

the diversity in eyespot colour composition, across an extensive phylogeny of

Mycalesina butterflies. Results are largely consistent with those from B. anynana, with

the notable exception of a diverse clade on Madagascar. Thus, whereas each species

generally has eyespots of similar colour-composition, consistent with a major role for

developmental bias in shaping evolutionary diversification, most Malagasy Heteropsis

species have gained independent control of eyespot colour-composition enabling a

wider exploration of morphospace. We then investigated the formation of this novel

phenotype by manipulating eyespot development in early pupal wings of H. iboina. The

results show that the bias has been broken through a novel difference in the response

of different areas of wing tissue to a conserved eyespot patterning signal. Our study

demonstrates how development can potentially constrain the evolutionary

independence of traits, but that at a macro evolutionary level the constraint can be

released by an innovative developmental change allowing rapid morphological change

and facilitating evolutionary diversification. Future directions include more detailed

studies of how the bias was broken in the early Heteropsis radiation, and whether the

bias found in most Mycalesina might be adaptive.

38

Fighting about frequency

Karen Kovaka

Virginia Tech, USA

Many controversies in biology, including many of those at the heart of the Extended

Evolutionary Synthesis, have two distinctive features. First, they are not controversies

about whether some hypothesis is right or wrong. Rather, they are controversies about

how often a certain pattern or process occurs. Second, these controversies are rarely

resolved in favor of one side or another. They are more likely to fizzle out or be

abandoned. Examples of historical controversies that have these two features include

disputes about (a) punctuated equilibrium versus gradualism, (b) sympatric versus

allopatric speciation, (c) genetic drift versus natural selection, and (d) genetic versus

extra-genetic inheritance.

I provide an account of what justifies these kinds of controversies, and of how they

contribute to science. This is an important project, because many criticisms of the

Extended Evolutionary Synthesis rely on skepticism about the value of and justification

for such controversies.

More specifically, I argue for three claims:

1. There are serious questions about the value of controversies with the two

features I described above. How can we rationally justify scientific controversies that

rarely reach a resolution? Why should any scientist participate in such a controversy if

they can know in advance, with high probability, how the controversy will end?

2. The areas of research encompassed by the Extended Evolutionary Synthesis

include many controversies that have the two features I described above.

3. Despite the serious questions raised in (1), these kinds of controversies are

rationally justified. Engaging in these controversies improves the quality of the

explanations that biologists are able to give about the phenomena central to the

controversies."

39

On leaders and followers: the evolution of phenotypic plasticity can lead the

evolution of genetic evolvability more easily than vice versa

Miguel Brun-Usan, Alfredo Rago, Richard A Watson

University of Southampton, UK

In conventional evolutionary theory, phenotypic plasticity is often thought to be a

genetically encoded feature which can be evolved autonomously. However, since both

the effect of environmental conditions on phenotypes (the ExP map underlying

plasticity) and the effect of genetic mutations on phenotypes (the GxP map underlying

evolvability) depend on development, changes in one may affect the other.

But which is the leader and which is the follower? By means of computational

modelling, we show that it is much easier for the evolution of plasticity to lead the

evolution of genetic evolvability than vice versa. Two complementary reasons explain

this observed asymmetry: First, genetic and environmental inputs affect development

in different mechanistic ways, and thus the GxP and the ExP maps, although related, do

not exhibit identical dynamical nor variational properties. Second, the evolution of a

GxP map is more limited than the ExP map. In the GxP map, selection on phenotypes is

strong and selection on GxP maps is relatively weak. In contrast, the evolution of an ExP

map does not suffer these limitations: an individual can be selected according to its

ability to produce a distribution of phenotypes under different environmental

conditions.

We demonstrate that this asymmetry is consistent under different modelling strategies,

parameter combinations and experimental setups, thus suggesting that phenotype-first

scenarios may be the norm rather than the exception in biological evolution. Put in

simpler words: phenotypes are more often the leaders, and genotypes are more often

the followers."

40

Bird nests evolve flexibly in response to egg characteristics

Catherine Sheard, Sally E Street, Mary C Stoddard, Camille Troisi, Andrew Clark, Susan

D Healy, Kevin N Laland

University of St Andrews, UK

Cultural evolution allows humans to dramatically modify their own selective

environment (e.g. building shelter), a process known as ‘niche construction’. A classic

non-human example of niche construction is avian nest building, which recent evidence

in both the lab and the wild has shown can be socially transmitted. Here, we explore

the co-evolution of avian nest building and egg traits. We compiled a database of nest

structure and location for 8,231 species of birds, which we combined with a new

database of egg colouration for 4,762 species and previously-published information on

the egg shape of 1,400 species (Stoddard et al. 2017). Using Bayesian phylogenetic

comparative methods, we show that nest structure and location is correlated with egg

size, shape, colouration, and patterning. In many cases, however, it is the egg trait that

evolves first, followed by a corresponding change in the nest characteristic. On a global

scale, these egg traits are related to a variety of intrinsic and extrinsic factors. It has

been previously shown that egg size is tightly correlated with body mass and that egg

shape is related to adaptations for flight ability (Stoddard et al. 2017); we additionally

demonstrate that egg colour and patterning is correlated with environmental variables

and body mass, finding no support for traditional hypotheses related to brood

parasitism or sexual selection. Taken together, this evidence runs against the

conventional view of nests as highly conserved constraints on avian reproduction and

instead suggests that nest construction is a flexible behavioural trait that can quickly

respond to developmental sources of selection, including properties of the eggs.

Stoddard, M. C., E. H. Yong, D. Akkaynak, C. Sheard, J. A. Tobias, & L. Mahadevan. (2018)

Avian egg shape: Form, function, and evolution. Science 356: 1249-1257.

41

Adaptive evolution NOT driven by natural selection (?!?). On individual responses,

inclusive inheritance, and fleshing out the second term of that famous Price equation.

Pim Edelaar

University Pablo de Olavide, Spain

Evolutionary adaptation is typically accredited to natural selection only. However,

natural selection can only adapt populations, and by itself has little to offer to locally

maladapted individuals. Because of this, a large array of flexible individual responses to

environmental variation have evolved that can help individuals increase their fitness.

By means of a simple yet apparently comprehensive classification framework, I will

derive how these can be grouped into three distinct classes (as appears to be confirmed

from the social and economic sciences): adjustment of the phenotype (e.g. plasticity),

adjustment of the environment (e.g. niche construction in the strict sense), and

selection of the environment (e.g. habitat choice). I will then explore to what extent,

and under what conditions, these flexible responses can additionally and independently

drive adaptive evolution. I will do so by using the Price equation for evolutionary change

as a context, focussing on the rather neglected second term describing parent-offspring

resemblance, and touching upon discussions surrounding the Extended Evolutionary

Synthesis, the Extended Phenotype, and inclusive inheritance. I think this view can help

to shift our evolutionary thinking from an organism- and gene-centred position towards

that of the evolution of the organism-environment interaction via genetic and

alternative hereditary mean, while remaining consistent with basic evolutionary theory.

42

DNA methylation lends plasticity to the coloration of an African cichlid

Sebastian Alvarado

Queens College CUNY USA

Coloration is a salient trait across the animal kingdom that can allow an individual to

become cryptic, conspicuous, or social. While some developmental patterns in

pigmentation are static, others are dynamic to changes in their ambient environment.

For example, seasonal variation in the Arctic allows mammalian species to gain and lose

a white winter coat. Despite a great deal of study in developmental pigmentation

patterns, little is known about how environmental cues shape the molecular

mechanisms that allow an animal to change color. It stands to reason that the

underlying mechanisms of a plastic trait must also be plastic. One approach to

understanding these processes is through the lens of epigenetic modification and DNA

methylation. DNA methylation of cytosine residues in gene promoters is a reversible

modification that silences gene function in vertebrates. Since DNA methylation is

involved in programming various cellular functions, it is likely that it facilitates

molecular changes as pigment-bearing cells (chromatophores) change their

composition during animal color changes. In this study, we used an African cichlid model

system (Astatotilapia burtoni) with discrete reversible color morphs (blue and yellow)

to dissect the underlying molecular processes that lend plasticity to animal coloration.

We describe dynamic changes of DNA methylation within candidate regulators of

endothelin signaling to demonstrate control over the function of yellow

chromatophores. Our findings suggest that epigenetic processes such as DNA

methylation lend plasticity to coloration, which is an important hallmark driving

selection in Lake Tanganyika. Furthermore, since genetic diversity does not account for

the phenotypic diversity seen in Lake Tanganyika, we propose that DNA methylation

may contribute to the processes that have led to the adaptive radiation of cichlids in

East African Great Lakes.

43

Unexplored maternal effect mediators, thyroid hormones, in an eco-evolutionary

context

Suvi Ruuskanen, Bin-Yan Hsu, Tom Sarraude, Antoine Stier

University of Turku, Finland

Over the past decades, studies in maternal androgens and glucocorticoids have

demonstrated that maternal hormones are key mediators of maternal effects,

constituting an important source of transgenerational phenotypic variation.

Nevertheless, maternal thyroid hormones (THs), despite their well-known importance

on development in humans, have been largely neglected in ecological and evolutionary

context. We studied environmental and genetic variation of these maternal effect

mediators (biologically active T3, and prohormone T4), and associations between

maternal THs and offspring development, phenotypic variation and fitness via

experimental manipulation. Maternal yolk THs showed variation in relation to key

maternal extrinsic and intrinsic factors (temperature and laying order), and T3 was

moderately heritable. Experimental egg TH manipulations in five avian species revealed

that maternal yolk THs influence hatching success, growth, physiology (metabolic rates,

oxidative status) and aging markers, but the direction of the effects was highly

heterogeneous across species. Context-dependent effects may be one explanation for

such discrepancy, and is currently being tested. While no effect of elevated maternal

yolk THs on subadult survival was detected, effects on reproductive success have not

yet been assessed. However, many of the traits affected by maternal yolk THs are of

known importance to fitness, pending selection on these maternal effect mediators.

These results emphasize that maternal THs can be additional players of inclusive

inheritance and their role e.g. in adaptive developmental programming should be

further investigated.

44

Phenotypic plasticity facilitates colonization of savannas in Mycalesina butterflies

Sridhar Halali, Hisham Ali, Paul M Brakefield, Oskar Brattström


Phenotypic plasticity has been considered to facilitate persistence of populations in new

environments and the colonization of novel ecological niches. We test this hypothesis

using butterflies of the genus Bicyclus in which many species show adaptive

developmental plasticity with respect to seasonal changes. The rapid diversification of

Bicyclus during the Miocene period has been linked to colonisation of expanding

savannah grasslands. However, colonizing savannahs would likely have required a suite

of adaptations to cope with seasonally fluctuating resource availability compared to the

more stable ancestral forest habitats. Here we test two hypotheses: Firstly, savannah

species will show a higher degree of plasticity than forest species. Secondly, plasticity

may have facilitated the colonization of novel savannah habitats.

We derived a plasticity index for 100 Bicyclus species by measuring the differences in

eyespot size between wet and dry season morph in over 1000 museum specimens.

Furthermore, we quantified the niche of each species using Environmental Niche

Modelling based on museum location data and seven climate variables.

The plasticity index shows a moderate phylogenetic signal (Pagel’s λ, 0.64) following an

Ornstein-Uhlenbeck model of trait evolution. Savannah species exhibited a higher

degree of plasticity, but forest species had higher variance (PGLS, p<0.01). Three

seasonally linked climatic variables explained 70% of the variation, and correlated

significantly with the plasticity index (PGLS, p<0.01).

Phenotypic plasticity can modulate optimal season-specific trait expression in these

butterflies which is necessary for survival, especially in the savannah. Also, the plasticity

index is expected to evolve in response to habitat-specific adaptive optima. An

observed higher variance among forest species is partly explained by phylogenetic

inertia coupled with low cue reliability in more stable environments. Highly plastic

species generally occupy extreme niches suggesting that plasticity may have been an

important trait in facilitating colonization of savannas and facilitated diversification of

Bicyclus.

45

Does thermal plasticity align with local adaptation? – An interspecific comparison of

wing shape and size in sepsid flies

Patrick T Rohner, Wolf U Blanckenhorn, David Berger

University of Zurich, Switzerland

Although genetic and plastic responses are sometimes considered as unrelated

processes, their phenotypic effects may often align because genetic adaptation is

expected to mirror phenotypic plasticity if adaptive, but to run counter to it when

maladaptive. The magnitude and direction of this alignment has further consequences

for both the tempo and mode of adaptation. To better understand the interplay

between phenotypic plasticity and genetic change in mediating adaptive phenotypic

variation to climate variability, we quantified genetic latitudinal variation and thermal

plasticity in wing loading and wing shape in two closely related and widespread sepsid

flies. Common garden rearing of 16 geographical populations reared across multiple

temperatures revealed that wing loading decreases with latitude in both species.

Having evolved convergently, this pattern could be driven by selection for increased

dispersal capacity in the cold. However, although allometry, sexual dimorphism,

thermal plasticity and latitudinal differentiation in wing shape all show similar patterns

in the two species, the relationship between the plastic and genetic response differed

between them. While latitudinal differentiation (south to north) mirrored thermal

plasticity (hot to cold) in Sepsis punctum, there was no relationship in Sepsis fulgens.

While this suggests that thermal plasticity in wing shape may have contributed to

mediate local adaptation in S. punctum, it also demonstrates that genetic wing shape

differentiation and its relation to thermal plasticity may be complex and idiosyncratic,

even among ecologically similar and closely related species. Hence, genetic responses

can, but do not necessarily, align with phenotypic plasticity induced by changing

environmental selection pressures. We discuss the potential role of plasticity in biasing

adaptive genetic change and illustrate the usefulness of comparing replicated genetic

and plastic responses of complex phenotypes in response to analogous selection

pressures in revealing adaptive trait variation.

46

Macroevolutionary dynamics of phenological plasticity

Stephen De Lisle, Erik Svensson

Lund University, Sweden

Theory predicts that adaptive plasticity in fitness-related traits may play a key role in

establishment in novel environments, persistence in changing environments, and

allopatric speciation. Yet testing these hypotheses is difficult, especially at the

macroevolutionary level, due to the inherent difficulty of measuring plasticity. We

exploit recent methodical advances to estimate the strength of plasticity and the

environmental dependence of selection, the two key parameters linking plasticity and

local adaptation, from observational data from a largescale database of over one million

individual records of adult damselflies and dragonflies and corresponding spring

temperatures. Our aim is to elucidate the role that temperature-induced plasticity in

timing of metamorphosis (phenological plasticity), a key life history transition, plays in

macroevolutionary diversification. We first provide a validation of the methods using

an independent dataset of phenotype and fitness from a single damselfly species,

showing a striking correspondence between the observational estimate of the

environmental dependence of selection and the direct estimate. We then use the

observational approach to test two theoretically-motivated predictions: 1) the strength

of plasticity should coevolve with the environmental dependency of selection and 2)

the contribution of plasticity to local adaptation should be highest in extreme/recently

colonized environments. Our analysis of over one million records from 49 species

supports both predictions. First, we find correlated evolution of the strength of

plasticity and the environmental dependence of selection. Second, we show that the

contribution of phenotypic plasticity to within-species local adaptation increases during

the recent, post-glaciation colonization of Northern regions. Our results suggest that

phonological plasticity may have played an important role in temperate diversification

in these insects.

47

Developmental regulation and evolution of nutrition-responsive growth:

transcriptomic underpinnings of plasticity in horned beetles

Sofia Casasa, Eduardo E Zattara, Armin P Moczek


Plastic responses to nutritional variation are ubiquitous features of developmental

systems, yet our understanding of the mechanisms that regulate nutritional plasticity

as well as bias its diversification is limited. We used comparative RNAseq analyses of

Onthophagus horned beetles to better understand the transcriptomic mechanisms

underlying the development and evolution of nutrition-sensitive plasticity of horns,

male-biased weapons used in competition over mating opportunities. We investigated

three Onthophagus species whose nutritional plasticity in horn formation ranged from

the ancestral, moderately polyphenic state to the derived exaggerated polyphenic

state, or alternatively, the secondary loss of nutrition responsiveness. We find that (i)

the regulation of polyphenic development is partly shared across species regardless of

the degree of nutrition-sensitivity, (ii) that diverse genes have been recruited into the

evolution, and in particular the exaggeration, of nutrition-responsiveness, and (iii) that

secondary loss of plasticity is paralleled by extensive loss of gene expression plasticity.

Our results suggest that nutrition-mediated plasticity in morphology is underlain by a

corresponding genome-wide magnitude of gene expression plasticity, and that both

novel as well as pre-existing gene regulatory networks contribute to the elaboration of

nutrition responsive development. By utilizing a comparative, unbiased, genome-wide

assessment our work provides comprehensive insights into the regulation and evolution

of nutrition-responsive development.

48

Involvement of maternal yolk thyroid hormones in the evolution of life history traits

in birds: a comparative analysis

Bin-Yan Hsu, Veli-Matti Pakanen, Tapio Eeva, Blandine Doligez, Ton GG Groothuis, Erkki

Korpimäki, Toni Laaksonen, Asmoro Lelono, Pat Monaghan, Tom Sarraude, Barbara

Tschirren, Robert L Thomson, Jere Tolvanen, Rodrigo A Vásquez, Suvi Ruuskanen

University of Turku, Finland

As mediators of prenatal maternal effects, maternal hormones can bring about

additional influence on evolutionary processes above and beyond genomic inheritance.

Maternal thyroid hormones (THs), well-known for the essentiality on embryonic brain

development, may harbour great evolutionary significance but our understanding to

this is extremely limited. Lately, a few experimental studies in birds demonstrated that

elevated maternal yolk THs within the natural variation influenced offspring

development and growth. However, these studies showed striking differences between

species, despite the essentially similar experimental designs. This thus calls for

comparative studies to investigate the evolutionary relationship between maternal THs

and life histories. In this study, we analysed maternal yolk THs from un-incubated eggs

from 34 bird species across 17 families and 6 orders, representing a wide range of body

size and life histories. Our phylogenetic comparative analysis suggested clear links

between maternal yolk THs across a few main life-history continuums. First, migratory

species deposited higher concentrations of THs in the egg yolks than residential species.

Because of the regulatory roles of THs in seasonal activities, this association suggests

the involvement of maternal THs in the evolution of migration or the circannual rhythm.

Second, precocial species deposit larger amounts of T3, the bioactive TH, in the egg

yolks than altricial species after controlling for species’ body masses. As the

development of thyroid gland starts much earlier in precocial species than in altricial

species, this association suggests the importance of maternal T3 to support the earlier

development of the hypothalamic-pituitary-thyroid axis. In conclusion, our data

suggested important involvement of maternal THs in the evolution of life histories. The

patterns we identified should provide important basis for further experimental work.

49

Phenotypic divergence across spadefoot toad species mirrors within-species

consequences and mechanisms of developmental plasticity

Christoph Liedtke, Ivan Gomez-Mestre

Doñana Biological Station, CSIC, Spain

Phenotypic divergence across derived lineages sometimes mirrors phenotypic plasticity

in the common ancestor, suggesting that environmentally induced phenotypic changes

may precede changes in genotype (i.e. ‘flexible stem hypothesis’) that can evolve into

species divergence through genetic accommodation. Pelobatoids, the spadefoot toads,

stand out as a good system for studying the role of plasticity in evolution because this

group shows extensive developmental rate variation within and between species with

predictable consequences on adult phenotype. Whereas Pelobates cultripes has

retained the highly plastic developmental rate plesiomorphic of this group, this

plasticity has been greatly reduced in Scaphiopus couchii as a consequence of

adaptation to ephemeral breeding ponds. As a result of this adaptive process

Scaphiopus couchii now presents several features that resemble an accelerated version

of Pelobates cultripes, from overall morphology to fat storages to metabolic rate and

endocrine regulation of development. To understand how the transcriptional

regulation may have evolved between these divergent species, we have also

characterized the gene regulatory network underlying developmental rate in both P.

cultripes and S. couchii. We have investigated whether the same underlying

transcriptional profile is conserved across species, and whether it varies between

species in its environmental sensitivity. Spadefoot toads provide a good example of how

canalization of ancestrally plastic traits can contribute to adaptive divergence and

evolutionary novelty.

50

What the history of biology tells us about the Extended Evolutionary Synthesis

Jan Baedke1, Daniel J Nicholson2

1Ruhr University Bochum, Germany; 2Konrad Lorenz Institute for Evolution and Cognition

Research, Austria

Recently, there has been a growing number of calls for expanding evolutionary theory.

Advocates of a so-called Extended Evolutionary Synthesis (EES) have argued that

phenomena studied in evo-devo, epigenetics, and niche construction theory should be

more seriously integrated into standard evolutionary theory. This includes the

conviction that evolutionary investigations should focus less on genes and more on

organisms and their reciprocal interactions with their environment. Unfortunately, this

recent debate lacks a historical understanding of what kind of expansion the EES

actually is (or could be) and on which theoretical traditions it draws upon. So far,

attempts to historically contextualize the EES have focused primarily on showing how it

differs from the Modern Synthesis (MS). We show that this historical perspective is

limited. We argue that while the EES introduces a battery of concepts, theories, and

explanatory standards to evolutionary biology compared to the MS, many of these are

actually quite old. In fact, the theoretical core of the EES is not ‘post-MS’ but rather ‘pre-

MS’. That is to say, the EES reintroduces claims into biology regarding the relationship

between genes, organisms, and their environment that were already clearly articulated

and vigorously defended by early twentieth-century biologists, especially organicists,

dialectical materialists and (German) holistic biologists. We discuss six recurring

theoretical claims: (1) ‘genes neither control nor determine development’; (2) ‘genes

are not the exclusive repositories of inheritance’; (3) ‘organisms act as agents in their

environments’; (4) ‘organisms and environments reciprocally constitute each other’; (5)

‘organisms extend into one another’; and (6) ‘organisms form together new collective

individuals’. If none of these claims are new, then why did they not become

incorporated into evolutionary theory much earlier? In the final part of our talk we shall

attempt to answer this crucial question.

51

Towards a novel unifying theory of the genotype concept

David Ricote Hernández, Ignacio Maeso

Centro Andaluz de Biología del Desarrollo, CSIC, Universidad Pablo de Olavide, Spain

Understanding the Genotype-Phenotype (G-P) interdependence is fundamental for all

biological fields. Surprisingly, discussions about the limits of the G-P dualism have been

sidestepped by mainstream scientific discussions, despite the fact that both

experimental practice (from mendelian genetics to -omics) and theoretical background

(from phenomenology to information theory) have changed profoundly since

Johannsen first coined the term genotype in 1911. Indeed, multiple unresolved

contradictions between the original G-P formulation and its current use are showing

that explicit theoretical reevaluations are urgently needed. In particular, the so-called

extragenetic inheritance phenomena have not been possible to accommodate to the

genotype concept due to the current lack of a neat distinction between genotype and

genome, excluding inherited epigenetic configurations as fundamental and integral

parts of the genotypical identity of organisms.

Here, we provide a novel proposal towards a unifying theory of genotype based on 4

objective and experimentally verifiable criteria (self-templating, translatability,

plasticity and inheritability) that are able to account for all potential sources of

biological inheritance as genotypic constitutions, from molecular structures to social

behaviour. Our proposal follows a different path than previous related frameworks

(such as the replicator concept) because it does not rely on information to delineate the

boundary between genotype and phenotype, but on the evolvability conferred by a

growing number of nested and interrelated genotypical systems. By doing so, we

provide new solutions to several long-standing problems, from the periodic resurgences

of lamarckian phantoms to controversies on biological levels and units of selection.

52

Rethinking cybernetics in contemporary theoretical biology

Flavia Fabris

Egenis, the Centre for the Study of Life Sciences, University of Exeter, UK

In recent years, the contributions of cybernetics to the development of evolutionary

developmental (evo-devo) biology have increasingly been recognised. The particular

theories and models developed during the flourishing of cybernetics in the early 20th

century laid the foundation for the systems approach, which is nowadays widely and

fruitfully employed in molecular biology, genetics, genomics, immunology,

developmental biology, and ecology. Nevertheless, no philosopher or biologist seems

to know what cybernetics is, and often what they think they know they dislike:

cybernetics is often identified with a reductive ‘machine conception’ of the organism

and an engineering view of biology. However, once we understand what cybernetics is

really about, we see such conceptions are mistaken and moreover that a cybernetic

perspective can shed significant light on major discussions in current biology and its

philosophy: in particular, on the fate of the Modern Synthesis in light of later

developments in biology, the purpose and nature of evolutionary developmental

biology, and disputes between those who emphasize a mechanistic conception of

biology and ‘processualists’. Thus, my current research has two objectives: the first is

to clarify the relationship between cybernetics and reductionism, and the second is to

demonstrate the relevance of cybernetics to evo-devo. To accomplish the first

objective, I will provide positive arguments for the thesis that, in contrast to the

predominant view, cybernetic explanations within biology, when properly understood,

are non-reductionistic, and do not have, at their core, any heavyweight metaphysical

commitment to the mechanistic nature of life. To accomplish the second objective, I

will disentangle the nature of cybernetics and reappraise its history in order to show

how it offers new tools for approaching well-known neo-Darwinian controversies that

have emerged in recent years.

53

The Extended Evolutionary Synthesis debate: some epistemological and sociological

dimensions

Alejandro Fábregas-Tejeda, Francisco Vergara-Silva

Universidad Nacional Autónoma de México, Mexico

For the proponents of the Extended Evolutionary Synthesis (EES), the narrow and ‘gene-

centric’ stance of the Standard Evolutionary Theory (SET) fails to capture the full gamut

of causal processes entangled in the evolution of biological diversity (in particular,

developmental bias, niche construction, developmental plasticity, and extra-genetic

inheritance). In that sense, the EES denounces an incomplete ontology of evolutionary

processes in the SET. The EES, as a conceptual framework that emphasizes organismal

causes of development, inheritance and differential fitness, the role of constructive

processes in development and evolution, and reciprocal representations of causation,

delivers an extended ontology of evolutionary processes, including those that generate

novel variants, bias selection, modify the frequency of heritable variation, and

contribute to inclusive transgenerational inheritance. Here, we discuss two far-reaching

dimensions of the EES debate: i) epistemological: by focusing on the explanatory goals

of this framework, (purported) interdisciplinary integration, the conceptualization of

the ‘organism-environment’ relationship, and tensions subtending the notions of

‘developmental system’, ‘organismal agency’ and ‘developmental niche(s)’; ii)

sociological: by analyzing discursive strategies involved in the disagreement between

the defenders of both the SET and the EES, the dispute for ‘symbolic and scientific

capital’ (sensu Bourdieu), ethical-political implications of this new view of evolution,

and other issues that can be unmasked by STS-studies and ideas coming from recent

sociology of science. Finally, we argue that a practice-oriented analysis of the EES

debate may bridge some of the epistemological and sociological dimensions previously

discussed.

54

Beyond standard views of reproduction: pregnant females as individuals and their

role in evolution

Arantza Etxeberria, Laura Nuño de la Rosa, Mihaela Pavlicev

Complutense University of Madrid, Spain

While reproduction is generally agreed to be a crucial feature of life on Earth, its roles

in evolution have been neglected, either by considering it secondary to self-

maintenance, or by reducing it to replication and copy making. In the latter case, the

highly diverse modes of reproduction explored in evolution are examined as

reproductive strategies with a single purpose: to maximize fitness, without considering

the effects of diverse reproductive modes, for example, on relations between biological

entities. Recently, there has been an increasing call to consider the material processes

involved in reproduction, and evo-devo seems to be the natural disciplinary candidate

for integration of reproduction and development into the structure of evolutionary

theory. The theoretical implications of an “evo-devo of reproduction” have remained

largely unexplored so far, also due to its focus on the evolution of body parts and their

interactions. In this presentation, we will reflect on the crucial implications of recent

research in evolution of therian reproduction, particularly of the origins of eutherian

pregnancy, for the status of the pregnant female. This problem, namely if the pregnant

female is a carrier of fully-fledged developing individuals or a single individual by itself,

is becoming a focus of debate in the philosophical literature. Our goals will be to 1)

question some common assumptions about biological reproduction and individuality

for evolution; 2) explore, in the light of recent work in evo-devo, how mammalian

reproduction has evolved as a merging of female and egg that involves major

innovations at different (morphological, physiological, developmental, genetic) scales

of organization; 3) consider what kind of individuals are involved in pregnancy from the

physiological, evolutionary, and interactive perspective; 4) propose that, from a

reproductive perspective, the pregnant female as a developmental stage can be

considered tantamount to a evolutionarily novel kind of temporary individual.

55

Evolution of multicellularity: cheating done right

Walter Veit

University of Bristol, UK

For decades Darwinian processes were framed in the form of the Lewontin conditions:

reproduction, variation and differential reproductive success were taken to be sufficient

and necessary. Since Buss (1987) and the work of Maynard Smith and Szathmáry (1995)

biologists were eager to explain the major transitions from individuals to groups

forming new individuals subject to Darwinian mechanisms themselves. Explanations

that seek to explain the emergence of a new level of selection, however, cannot employ

properties that would already have to exist on that level for selection to take place.

Recently, Hammerschmidt et al. (2014) provided a ‘bottom-up’ experiment

corroborating much of the theoretical work Paul Rainey has done since 2003 on how

cheats can play an important role in the emergence of new Darwinian individuals on a

multicellular level. The aims of this paper are twofold. First, I argue for a conceptual

shift in perspective from seeing cheats as (i) a ‘problem’ that needs to be solved for

multi-cellularity to evolve to (ii) the very ‘key’ for the evolution of multicellularity.

Secondly, I illustrate the consequence of this shift for both theoretical and experimental

work, arguing for a more prominent role of ecology and the multi-level selection

framework within the debate then they currently occupy.

56

The explanatory robustness of developmental types: a propensity view

Cristina Villegas

Complutense University of Madrid, Spain

In this talk I want to provide a philosophical frame for understanding the explanatory

role of the properties of developmental systems explored in the field of evo-devo. A

propensity interpretation of variability and evolvability is first introduced. These

properties will be claimed to be probabilistic dispositions responsible for the variational

patterns represented in evo-devo models. Following Roberta Millstein’s suggestion, in

the context of the nature of fitness discussion, that a long-run propensity interpretation

of biological probabilities is best understood as a type propensity interpretation, I will

defend that the explanatory power of such evo-devo properties partially relies on their

reestablishment of a certain kind of typological thinking without necessitating the –

always controversial– reification of types. I will then argue that this feature provides

evo-devo explanatory models with greater counterfactual robustness, enabling them to

address how-possibly questions about the generation of variation and innovations

more satisfactorily than populational explanations.

57

The cultural evolution of cultural evolution

Jonathan Birch, Cecilia Heyes

London School of Economics and Political Science, UK

Human cultures are not the only cultures. Other primates, as well as cetaceans and

birds, pass learned skills down the generations. But human cultural evolution is

different. It is cumulative: small improvements to skills build up. Moreover, the

accumulation is fast: improvements spread rapidly, no longer tied to the timescale of

biological generations. How does fast cumulative cultural evolution work? Where did it

come from? And why is it the sole preserve of humans?

One approach to these puzzles appeals to genetic adaptations that only humans

possess. Heyes (Cognitive Gadgets, HUP, 2018) develops a novel approach on which the

mechanisms of cultural learning are products of cultural, not genetic, selection. Here

we turn this explanatory strategy towards the process of “cultural selection"" itself. In

short, we propose that cultural selection itself evolves culturally.

To motivate this idea, we highlight a crucial distinction between two types of cultural

selection: CS1 and CS2 (Birch, The Philosophy of Social Evolution, OUP, 2017). CS1 is

variation in biological fitness due to culturally transmitted variation, whereas CS2 is

variation in cultural fitness (roughly, the number of learners a model recruits) due to

culturally transmitted variation. We explain how ""cultural fitness"" should be defined,

and we distinguish two cognitive “grades” of CS2, depending on whether model

selection by learners is based on attentional or metacognitive processes.

This allows us to pose the question: which type of cultural selection enables fast

cumulative culture, and how did this type of process originate? We hypothesise that

metacognitive CS2 (and not merely attentional CS2) is essential for cumulative culture.

We further hypothesise that metacognitive CS2 was assembled by simpler, earlier

processes of CS1. We draw on evidence from cognitive science, and especially

developmental psychology, to support our hypotheses."

58

Evolution by natural induction

Richard A Watson


It is widely assumed that natural selection is the only possible source of adaptation

occurring spontaneously in natural systems. This assumption short-circuits any debate

over the significance of other processes in evolutionary change, e.g. the ‘active roles of

phenotypes’. Induction, characteristic of learning systems, can also occur

spontaneously in suitable dynamical systems. For example, Hebbian learning, sufficient

for many non-trivial adaptive behaviours, occurs spontaneously in a physical system of

particles connected by springs if the springs are slightly plastic (imperfectly elastic).

When the system is repeatedly exposed to external forcing, its ability to find low-energy

state configurations under future forcing is increased - not because it was designed or

evolved to do so, but because of the direct action of differential forcing on its internal

organisation. Biological networks with evolving components are capable of such

inductive learning. This involves the action of natural selection only on individual links

in the network using only local selective feedback (equivalent to imperfect elasticity).

The result is that evolving systems, e.g. gene-regulation networks, can increase their

evolvability without being selected for this function. Interestingly, evolutionary models

cannot exhibit induction unless they include active phenotypes (e.g. dynamical

developmental processes). We thus conclude that evolution in natural systems

proceeds by a combination of natural selection and natural induction, that natural

induction is an adaptive mechanism in its own right (not a product of natural selection),

that the active roles of phenotypes are necessary for evolving systems to exhibit natural

induction, and that the adaptation provided by natural selection alone is significantly

limited in comparison. These conclusions support the view that the active roles of

phenotypes are important to natural evolution, not because they are proximal details,

but because they enable access to a different adaptive mechanism that is absent in the

standard model.

59

Generalized pathways to specific outcomes: stress links maternal environment and

adaptive offspring trait variation

Ahva Potticary, Renee Duckworth


The evolution of an adaptive maternal effect requires complex coordination of a

maternal response to environmental cues, coevolution of maternal physiology and

offspring embryonic development, and subsequent matching of adult offspring

phenotype to a future environmental context. How can such complex multi-

generational and multi-environmental coordination evolve? One possibility is that

evolution of maternal effects does not require de novo evolution of maternal

physiology, but instead coopts pre-existing pathways to link ecological cues to aspects

of maternal physiology that are primed to influence offspring development. An obvious

candidate for such cooption are maternal stress pathways, as they are highly conserved

across taxa and reactive to environmental context, providing a generalized cue of

environmental quality. We test this idea using a known adaptive maternal effect in

western bluebirds, where competition over a limited breeding resource (cavities in

trees) results in mothers influencing offspring dispersal strategies. We combine 17 years

of data on natural variation in female corticosterone levels and environmental

conditions with experimental manipulation of breeding resources to determine

whether multiple cues influence maternal stress, and whether maternal stress, in turn,

induces the maternal effect in offspring. We found that females had higher levels of

baseline corticosterone when breeding on territories that had either heightened

competition for nest cavities or more difficult foraging conditions during inclement

weather. Moreover, we found that irrespective of the environmental cue, the maternal

effect was associated with stress females experienced during oogenesis: females with

lower corticosterone levels produced more philopatric sons compared to females with

higher corticosterone levels. These results suggest that stress pathways are a general

link between ecological cues and offspring development and may foster the evolution

of maternal effects by enabling species-specific stressors to produce adaptive variation

in the offspring generation.

60

Evolution of migratory behavior by barnacle geese: the roles of experience, memory

and social learning

Thomas Oudman, Ingunn Tombre, Paul Shimmings, Graeme Ruxton, Jouke Prop

NIOZ Netherlands Institute for Sea Research, The Netherlands; University of St Andrews,

UK

Many migratory animal populations adjust their migratory decisions to changes in

spring phenology – some even stop migrating altogether. In many species, individuals

may learn their migratory strategy by following others. This opens up the possibility for

the cultural inheritance of strategies that are adjusted in response to individual

experiences.

Forty years of observations at multiple locations show strong changes in the use of

staging sites by barnacle geese along the Norwegian coast, on their way from the UK to

the breeding grounds on Svalbard. Barnacle geese are highly social animals, and the

choice of staging site strongly affects their reproductive success. This choice is not

straightforward. A reconstruction of foraging conditions at the different staging sites

from daily local temperature and radiation data showed that annual variation was high,

and that each staging site showed a different trend over the last forty years.

To understand how barnacle geese navigate this dynamic environment, we designed a

set of individual-based simulation models, each of them assuming a different set of

processes to underlie the choice of staging site. These include group size, reliance on

individual memory and exploration, and different social copying rules (e.g. based on

chance, relatedness, previous success or experience). This allowed us to explore how

each of these processes and their interactions would affect the population response to

the observed environmental variation. By comparing these simulated responses to the

actual population trends, we infer which of these processes are most likely to underlie

the migratory behavior of barnacle geese. We find that both social and individual

learning required assumptions for the simulations to resemble the actual population

trends. The simulations also point out additional types of field observations that will

help to discern the processes by which migratory behavior adjusts to changing

environments in other species.

61

Theory-based ecology and niche construction

Liz Pásztor

Biological Institute, Eötvös Loránd University, Hungary

Niche construction has been discussed in several ecological contexts. The theory of

niche construction differs from the standard theory of evolution in focusing on the

environmental effects of the organisms and on the selection processes induced by these

modifications. It can be linked easily with the theory of ecosystem engineering by

considering the selective effects of the nutrient cycles modified by the organisms. This

was modeled directly in a framework called „ecological niche construction”. The

selective effects of niche construction can also be integrated to community ecology by

considering selection in the context of species interactions.

However, these achievements have not revealed the common roots of the theory of

niche construction and the Darwinian theory of evolution and ecology. A theory-based

ecology (TBE) has been developed, which is founded on Darwinian principles and

defines ecological niches deductively. Ecological niches are described in terms of

population regulation including both positive and negative interactions among

populations. As TBE considers the environmental factors regulating population growth

explicitly the impact of population growth on these factors and reciprocal causation are

inescapable elements of this theory. Thus, evolution by niche construction follows from

first principles. This will be explained and illustrated in this talk.

62

The Hot Spring Hypothesis for the origin of life: implications for niche construction

theory in the first major transition in evolution from non-life to life

Bruce Damer, David Deamer, John Odling-Smee

The Biota Institute, USA; University of California, Santa Cruz, USA

Until recently the origin of life was thought to have occurred deep in the oceans at

hydrothermal vents as some energy sources and building blocks are available there.

However, chemical, thermodynamic and combinatorial barriers have led to an

alternative hypothesis situated in an updated version of Charles Darwin’s “warm little

pond”: fresh water volcanic hot spring pools that undergo cycles of hydration and

dehydration. The hypothesis involves construction of proto-niches that self-assemble

from membranous structures and guide chemical reactions related to synthesis and

evolution of polymer systems. Combinatorial selection amplifies functional polymers

within protocells ultimately incorporating heritable genetic memory. This hypothesis

has been tested experimentally in laboratory and volcanic hot spring conditions

suggesting that the first major transition from non-living compounds to living molecular

systems will be characterized by:

1. A cycling natural “engine” utilizing crowding to shape probabilities analogous to

loading the dice, a network effect of interacting polymers in protocell aggregates, and

Darwinian selection arising through an emergent memory system;

2. Communities of dividing living cells emerging from earlier self-assembling transitory

protocells;

3. The first genotypes arising through combinatorial selection of proto-phenotypes;

4. Vertical descent in genetic evolution originating from horizontal network sharing of

the products of chemical evolution.

The hot spring hypothesis not only provides an experimentally-accessible predictive

chemical framework for the transition to life but also a simplified exploratory laboratory

which could provide insights into the fundamental operating principles of evolution

itself. An international community has assembled around the testing of this hypothesis

and we will present compelling empirical results by the date of the meeting.

63

Developmental bias and morphological convergence in Anolis lizards

Nathalie Feiner, Illiam Jackson, Tobias Uller


Instances of convergent evolution provide insights into the processes that give

evolution directionality. While convergent phenotypes are usually attributed to

similarity in selective pressure, it remains unexplored to what extent convergence is

shaped by developmental biases. Yet, the fact that different morphologies can be

functionally equivalent (many-to-one mapping of phenotype-to-fitness) suggests that

morphological convergence is partly caused by biases in how development responds to

genetic or environmental perturbation.

After colonizing the four main islands of the Greater Antilles, Anolis lizards have

independently and repeatedly evolved into six ecomorphs that are well adapted to

running and climbing in certain microhabitats (e.g. grass, twigs, tree trunks or tree

crowns). In contrast, Lesser Antilles and mainland Anolis species do not obviously fall

into ecomorph categories, despite that they are typically found using similar habitat.

Using a morphological dataset (µCT-scans) of 350 Anolis species, we quantified the

extent of divergence and convergence in the relative size and shape of limbs, pelvic and

pectoral girdles between independent radiations of Anolis species. We quantified if

morphological divergence between species tends to fall along the same axis that

describes maximum variation within species. Furthermore, we tested the prediction

that adaptive divergence and convergence can be directed by developmental plasticity.

In an experimental setting with twelve species raised either in a ‘running’ or ‘climbing’

habitat, we established to what extent the relative size and shape of limbs and girdles

respond to mechanical stress induced by locomotion. These data allow us to assess to

what extent the patterns of adaptive divergence and convergence in morphology are

channelled by the process of development.

64

Measuring developmental bias: quantitative genetics meets developmental biology

Lisandro Milocco, Isaac Salazar-Ciudad


The patterns of inheritance of complex traits such as shape can be approximated by

quantitative genetics models. The models from this field have the core implicit

assumption that all phenotypic variation is symmetric, arising with equal likelihood.

There is however evidence that this assumption does not hold in many cases, a

phenomenon which has been called developmental bias. The term bias implies that

some phenotypic variation is more or less likely to arise, compared to a base

expectation, due to the dynamics of the developmental system involved in producing

that variation. The importance of this developmental bias in determining the direction

of evolution in the short term, however, remains heatedly debated.

In this work, we use the canonical equation from quantitative genetics, the multivariate

breeder's equation, as our base expectation for evolutionary change. We compare that

base expectation with the observed evolutionary change in an evolutionary simulation.

The latter combines a realistic genotype-phenotype map, from a tooth development

model, with a population-level model with sex, mutation and selection on the

phenotype. The deviations between the expected and the observed change are a

measure of the role that developmental dynamics play in constraining the directionality

of morphological evolutionary change. We found that the breeder’s equation often fails

to accurately predict evolutionary change because the assumptions of quantitative

genetics do not always hold for the morphological variation arising from development.

We quantify both the frequency and magnitude of these inaccurate predictions, which

can be as common as 31% for some traits and of large effect size. We found that these

prediction errors most commonly occurred for traits that are determined by the highly

nonlinear reaction-diffusion mechanism involved in tooth morphogenesis. These

results give a tangible measurement of the effect that developmental interactions have

in determining the direction of evolution in the short term.

65

The evolutionary potential of environmentally induced phenotypes

Reinder Radersma, Daniel WA Noble, Tobias Uller


Environmentally induced phenotypes have been proposed to initiate and direct

adaptive divergence. The evolutionary potential of environmentally induced

phenotypes depend in part on how well plastic responses are aligned with the additive

genetic variance and covariance among traits, which is captured by the genetic

covariance matrix (i.e., G matrix). Using meta-analyses, we show that the direction of

plastic responses to novel environments are generally aligned with the major axes of

standing genetic variation. Selection for or against environmentally induced

phenotypes is therefore likely to be effective. Furthermore, reciprocal transplant

studies reveal that evolutionary divergence tend to follow the direction of plasticity.

These results suggests that environmentally induced phenotypes may commonly

initiate adaptation to novel environments. However, the alignment between the

direction of plasticity and the main axis of genetic variation suggests that

developmental bias in response to environmental and genetic perturbation often may

be similar, making it challenging to distinguish between ‘plasticity-first’ vs ‘genetic

constraints’ scenarios of evolutionary divergence.

66

Developmental interactions established during speciation leave a lasting trace in

descendent populations

Laura Mears, Tom Ezard, Richard A Watson


Relationships among traits change the way that populations respond to selection. Traits

untethered by developmental interactions are free to evolve independently to match

the adaptive landscape. But, where developmental interactions exist, selection on one

trait elicits a response in others. To extract clues about how developmental

programming affects population divergence, we combine an adaptive dynamics

framework for modelling speciation with an individual-based model of evolvable

pleiotropic constraints. Using this system, we study how changes in development alter

the way that populations move over adaptive landscapes. Evolving independent traits

under strong selection results in among-individual variation orthogonal to the direction

of divergence. This orthogonal variation persists until populations reach static adaptive

peaks, at which time trait independence allows for rapid removal of non-adaptive

variation. But when evolvable interactions connect traits, among-individual variation

turns to align with the direction of divergence. We show that this directionality, and the

underlying developmental architecture, persist even after populations reach stable

adaptive peaks, preventing the loss of non-adaptive variation. Thus, the descendent

populations remain in alignment with one another, revealing the trajectory they

followed as the ancestral population diverged.

67

What kind of evolutionary novelty (if any) could be language?

Salvatore Ivan Amato, Giuliana Pulvirenti, Alessandra Falzone

University of Messina, Italy

The concepts of Novelty and Homology are as central in Evo-Devo approaches and in

the emergent Evolutionary Extended Synthesis as problematic and intertwined. These

could indeed be applied to different level of biological organization analysis; from

genetic and cellular to morphological as well as behavioral ones (Müller & Newman

2005; Minelli 2003). Studies at the phenotypic level have been able to identify several

generative mechanisms responsible of the emergence of evolutionary novelties

(genetic, epigenetic, behavioral, environmental). This theme has rightfully assumed a

great relevance within the biolinguistic area of research opening a theoretical debate

among Hauser-Chomsky-Fitch Vs Pinker-Jackendoff regarding the problem of the

emergence of the language faculty in our lineage: is it a unique feature of our human

nature or a mixture of several traits both shared and unique? Does it represent the

result of a gradual process of adaptation involving several components of the human

brain, or is it the result of a genetic mutation with a discontinuous outcome at the

phenotypic level? We think that the biolinguistic field is lacking of an adequate

problematization of the concept of novelty. This may be achieved widening the dialogue

with Evo-Devo/EES area of research, especially considering developmental plasticity

and niche construction processes (Pigliucci & Müller 2010; West-Eberhard 2003). An

analysis of the role that genetic, epigenetic and ecological factors may exert on the

formation of a phenotype, as well as the causal mechanism responsible of the

emergence of evolutionary novelties may foster a better characterization of the

linguistic phenotype and help elucidating the ways in which such phenotype comes to

be during the course of ontogenesis. This sort of investigation could also offer novel

insights about the evolutionary events and the possible mechanisms that have driven

its emergence during the phylogenesis.

68

It’s a horn, a wing, a helmet: the role of wing serial homologs in insect innovations

Yonggang Hu, David M Linz, Armin P Moczek


Understanding the origin of novel complex traits is a foundational challenge in

evolutionary biology. The most commonly used definition of novelty entails the absence

of homology to ancestral traits. This definition, however, is increasingly difficult to

reconcile with empirical findings across diverse taxa, which instead emphasize the role

of differential re-purposing of conserved developmental modules outside their

traditional developmental context as a dominant route to innovation. Yet how descent

with modification in developmental evolution may lead to morphological innovation

remains poorly understood, including the role of pre-existing gene networks in

enabling, biasing, or hindering such innovation. Here we employed RNA interference

mediated analysis of wing gene function with Hox-gene mediated transformations of

body segments to investigate the origin of the thoracic horns of scarabaeine beetles,

one of the most dramatic classes of secondary sexual traits in the animal kingdom. We

find (1) that thoracic horns derive from bilateral source tissues that fuse to form a single

medial outgrowth during development, (2) that diverse wing genes are functionally

required for instructing this process, (3) and that in the absence of Hox-input thoracic

horns transform into ectopic wings. Combined our results provide strong evidence for

the serial homology between thoracic horns and insects wings, and raise the possibility

that other insect innovations may similarly derive from wing serial homologs.

69

Beyond Darwin: how every individual adapts to its internal perturbations?

Yoav Soen

Weizmann Institute of Science, Israel

A critical gap in the theory of adaptation is the inability to explain how every individual

is able to cope with an unimaginably large number of epigenetic and symbiotic

variations, including many that are encountered in novel combinations and are unique

to that individual. Overcoming many of these novel, individual-specific variations

cannot be explained solely by natural selection, because the latter only accounts for

adaptation of the population as a whole. In other words, we understand how

populations adapt to changing environments, but we have no idea how an individual

adapts to stochastic internal changes of its own.

To address this gap, we proposed a non-traditional principle of individual-specific

adaptation, capable of extending the population-based adaptation by natural selection

to dynamic adaptations that are newly forming within every individual cell, animal and

plant. In some specific cases, acquired adaptations can even be inherited, thus enabling

progressive improvements in a few generations.

To investigate the validity of this Lamarckian theory, we established unique

experimental systems for testing the predictions of the theory in flies (D. melanogaster)

and in cultured cells. Both systems provide multiple lines of evidence supporting a

capacity to adapt by exploratory dynamics over time in an individual.

I will explain this concept of individual-specific adaptation and present substantial

experimental and theoretical evidence in support of this hypothesis.

70

How the beetle head got its horns: integrating a novel trait into a conserved trait

complex

David M Linz, Armin P Moczek


The origin of novel traits is among the most intriguing and enduring problems in

evolutionary biology. In particular, how novel complex traits are initiated and

incorporated within existing trait complexes without disrupting ancestral functions

remains poorly understood. The highly diversified horns of Onthophagus beetles

represent a recent evolutionary invention seamlessly integrated within the adult beetle

head, an ancient trait complex deeply conserved among insects. We sought to

investigate the developmental genetic network that instructs postembryonic head

formation in beetles, as well as identify the component parts that allowed horn

induction and positioning to become integrated into the dorsal head while maintaining

the integrity of head formation as a whole. To do so we executed a unique, fine-scale

RNAseq approach in both horned males and hornless females. This approach allowed

us to independently assess the transcriptional repertoires of six distinct head

compartments in both sexes. Contrasting these transcriptomes within and among sexes

then allowed us to (i) identify the transcriptional underpinnings of horn induction and

positioning in horn forming head regions as well as (ii) transcriptional changes occurring

in neighboring head regions putatively underlying compensatory, accommodating

development in the remainder of the head. Our results identify suites of genes critical

for patterning concise regions of the dorsal head and show that diverse embryonic

patterning genes have been repurposed to execute central roles in instructing the

formation of the postembryonic head, with some functions paralleling those executed

in the embryo, while others appear largely decoupled. Further, we identify points of

integration where re-use of pre-existing head patterning mechanisms enabled the

positioning of a novel trait – cephalic horns – within an otherwise conserved trait

complex – the dorsal head. We discuss our results by examining the roles of - and

interplay between - conservation and innovation in the origins of novelty and diversity.

71

Niche construction affects the consistency and strength of natural selection

Andrew Clark, Dominik Deffner, Kevin N Laland, John Odling-Smee, John Endler


Consideration of the properties of the sources of selection potentially helps biologists

to account for variation in selection. Yet, while investigations conducted over the last

40 years have generated many thousands of individual quantitative estimates of the

direction and strength of natural selection in the wild, associated analyses of such

environmental sources of selection and variation in selection are rare. Here we explore

how the variability of natural selection is affected by organisms that regulate

environmental variables through their activities (whether by constructing components

of their local environments such as nests, burrows, or pupal cases, or by choosing

suitable resources). Specifically, we test the prediction that organism-constructed

sources of selection that buffer environmental variation will result in reduced variation

in selection gradients, including reduced variation between (i) years (temporal

variation), and (ii) locations (spatial variation), and (iii) weaker directional selection,

relative to non-constructed sources. Using compiled datasets of 1045 temporally

replicated, 257 spatially replicated, and a pooled dataset of 1230 selection gradients,

we find compelling evidence for reduced temporal variation and weaker selection, in

response to constructed compared to non-constructed sources, and some evidence for

reduced spatial variation in selection. These findings, which remained robust to

alternative datasets, taxa, analytical methods, definitions of constructed/non-

constructed, and other tests of reliability, suggest that organism-manufactured or

chosen components of environments may have qualitatively different properties from

other environmental features.

72

Linking coral morphological traits and reef fish community structure

Luisa Fontoura, Stephanie D’agata, Kyle JA Zawada, Andrew H Baird, Mariana Álvarez-

Noriega, Rachael Woods, Michel Kulbicki, Maria Dornelas, Joshua S Madin, Joseph M

Maina, Elizabeth Madin

Macquarie University, Australia

The synchronized evolutionary expansion of small body size reef fishes with coral

complex forms in shallow waters suggests that branching corals might have had an

essential ecological role of providing shelter for fish to utilize. Losing structurally

complex coral forms can reduce shelter availability significantly for reef fishes, limiting

their ability to avoid predators and increasing competition for settlement area during

early-life stages. In this study, we aim to understand how changes in coral

morphological complexity, captured by coral convexity (a novel morphological trait),

affect the structure of reef fish assemblages. We use a multi-year dataset to track coral

community and reef fish assemblage structure, capturing a severe coral mortality event

that shifted the complexity of the coral community. We show that juvenile fishes, rather

than adults, are more responsive to shifts in morphological traits at the coral

community scale, a likely consequence of the higher vulnerability of juvenile reef fishes

due to competition for refuge in their early versus later life-stages. Changes in coral

morphological complexity explained 23% of juvenile reef fish decline, however the

magnitude of this effect varied by fish functional group. Although coral complexity

explained the decline of small body size juvenile fishes with restricted home ranges, we

found no relationship between changes in coral complexity and functional groups of

juvenile fish with large body size and wide home ranges. Differences in the responses

among fish functional groups due to loss of coral complexity suggest that structurally

complex coral species construct ecological niches that are occupied by vulnerable reef

fishes with small body size and restricted mobility. Importantly, our results shed light

on an evolutionary link between fish vulnerability and coral morphological traits.

73

Microbial community structure and function driven by infaunal niche construction in

intertidal sediments

Adam Wyness, Andrew Blight, Patricia Browne, Matthew Holden, David M Paterson


The Extended Evolutionary Synthesis theory predicts that, through their niche

construction, organisms greatly influence community structure, stability and

productivity, and co-direct evolutionary responses to selection in ecosystems (Odling-

Smee et al., 2003; Laland et al., 2015). The degree to which niche constructing activities

are complementary or conflicting impacts the magnitude and stability of biochemical

and physical gradients formed within sediment profiles. These gradients generated by

the ecosystem engineers are selective pressures to the microbial communities in

sediment. The aim of this work was to test how complementary and conflicting

ecosystem engineering impacts microbial community diversity and function and the

consequences on selected sediment functions such as surface stability, nutrient cycling

and primary production.

A well-established mesocosm system of intertidal sediments with treatments included

the presence of infauna (natural bioturbation with either one or both of Corophium

volutator, Nereis diversicolor), or manual turbation (proxy for bioturbation). Dissolved

oxygen and redox potential gradients were measured using microelectrodes. Microbial

community composition at the sediment surface and various depths was analysed using

high throughput 16S rRNA sequencing, and functional metagenomes predicted.

Evidence for the cooperative ecosystem engineering between the infauna of dissolved

oxygen and redox gradients compared to individual infauna treatments was observed,

and coincided with in shifts in diatom and prokaryote community composition. Surface

communities were compositionally and functionally different between infauna

treatments, with dominating influences observed in mixed treatments rather than an

intermediary response.

These results illustrate that niche construction is a powerful evolutionary force where

organisms modify their own and others’ selective environments. Furthermore,

observed differences in microbial community response to physical and bioturbation

hold significant inference for the impact of anthropogenic disturbance on

biogeochemical cycling by microbial communities. Now that community level effects

have been shown, this work will be taken further to examine ecological trajectories

among target bacterial populations.

74

Keeping close neighbours, corals are spatially clustered within and across species

Maria Dornelas, Shane Blowes, Tory Chase, Grace Frank, Mia Hoogenboom, Janine

Illian, Oscar Pizarro, Andrea Walker, Stefan Williams, Kyle Zawada, Joshua Madin


Spatial distributions of sessile niche constructing species determine coexistence

patterns. Major biodiversity models make two assumptions about spatial distributions:

that individual species are spatially clustered, and that the distributions of different

species are independent. These assumptions are common across all models that aim to

predict the global distribution of biodiversity. However, there is remarkably poor

empirical knowledge to support these assumptions. Moreover, to our knowledge these

patterns have never been quantified on reef corals, the niche constructors of the most

diverse marine ecosystem on the planet. We used AUV collected imagery to build 3D

topographic maps of a 7500 m2 of a reef at Lizard Island, Great Barrier Reef. These

models were annotated with the species identities and locations of over 11,000 coral

colonies, identified in situ by divers, to resolve the spatial distribution of the 120 species

in the surveyed area. We analysed these spatial distributions and quantified intra and

inter specific clustering. We show that across a range of scales, all but the rarest species

are spatially clustered with conspecifics, a finding that is consistent with model

assumptions. However, neighbouring species are remarkably diverse, and we find

evidence for interspecific clustering rather than independent distribution of different

species. We suggest that these patterns are likely to arise from niche constructing

mechanisms, such as clustering at the recruitment stage, when the physical structures

built by corals alter flow regimes in ways that facilitate settling of coral larvae.

75

Holed-up: nesting in a cavity as an evolutionary driver of parental care in fishes and

birds

Karina Vanadzina, Michael Webster, Kevin N Laland


Parental care is any form of behaviour that increases the survival rate or growth of a

parent’s offspring. Despite the wide taxonomic spread of parental care behaviours, we

know little about the evolutionary drivers of stable bonds between parents and their

young. The use of spatially restrictive breeding habitats does promote parent–offspring

association and could therefore contribute to the origin of care in clades such as ray-

finned fishes where more than 75% of families abandon their eggs shortly after

fertilisation. My analysis of reproductive behaviours in North American freshwater

fishes shows that cavity-spawners are 34 times faster at evolving parental care from the

ancestral state of no care than species that spawn in open waters. The evolutionary

impact exerted by the breeding habitat can also extend to clades where the majority of

species provide advanced care and raise their young in stable microhabitats such as

nests. For example, it has been hypothesised that hole-nesting passerines take care of

their young for longer periods and give rise to larger clutches when compared to birds

that nest in the open. This study is the first to explore the global distribution of parent–

offspring bonds in relation to breeding grounds in ray-finned fishes and passerine birds.

I use species-specific accounts of reproduction and the latest phylogenetic comparative

methods to analyse the co-evolution of parental care traits and breeding habitats. My

work fits within a framework of a larger project on niche construction that tests

whether artefact building and habitat choice generate consistent and predictable

selection pressures on a macro-evolutionary scale across a broad taxonomic spectrum

of animals.

76

Trait specialisation, innovation and the evolution of culture in changing environments

Dominik Deffner, Anne Kandler


Organisms often respond phenotypically to environmental challenges by innovating

and adopting novel behavioural variants. Behavioural (or 'cultural') variants are

alternative ways to solve adaptive problems, such as finding food or building shelter. In

unpredictable environments, individuals must both be able to adapt to current

conditions but also to cope with potential changes in these conditions, they must

‘hedge their evolutionary bets’ against the variability of the environment. Here, we

loosely apply this idea to the context of behavioural adaptation and develop an

evolutionary model where cultural variants differ in their level of specialization, i.e. the

range of environmental conditions in which they provide fitness benefits: generalist

refers to large ranges whereas specialist refers to small ranges. We use a Moran model

(with additional learning opportunities) and assume that each individual’s propensity

for asocial learning is genetically determined while the characteristics of cultural

variants are changed through asocial learning.

This model demonstrates that adjusting the level of specialisation allows organisms to

navigate the trade-off between fast initial adaptation and the potential for long-term

improvements. In situations with many learning opportunities, no adjustment of the

innovation rate (propensity to learn asocially) is required. Fast adaptation is guaranteed

by solely adjusting the level of specialisation of cultural variants. Few learning

opportunities, however, require both processes —innovation and trait specialisation —

to work hand in hand. To explore the effects of different modes of innovation, we

contrast independent invention and modification and show that a large proportion of

modifications improves both short- and long-term adaptation. Similarly, inaccuracies in

social learning provide another source of variant variation that facilitates adaptation

after an environmental change. However, unfaithful learning is detrimental to long-

term cumulative culture. Our results demonstrate that trait characteristics themselves

can play a major role in cultural dynamics and influence the evolution of learning

strategies.

77

A New Caledonian crow's-eye-view on how to develop probing intelligence: the

evolution of tool-use and tool-making as embodied, distributed and socially situated

perception action processes

Joanna E Fairlie1, Russell Gray2, Mei-Shin Wu2, Natalie Uomini2

1associated with the Max Planck Institute for the Science of Human History, Germany as

a guest researcher, 2Max Planck Institute for the Science of Human History, Germany

The behaviours of the tool-using Corvus moneduloides (New Caledonian crows or NCCs)

are often mentioned in support of existing theories about the evolution of Homo

sapiens cognition. Theories about their behaviours have been interpreted within

anthropological frameworks that have rarely considered modern evolutionary biology,

neuronal plasticity, ecological or developmental issues, or niche construction. Here we

discuss fresh cognitive models and behavioural approaches which allow us to consider

the NCCs in their own right as modern birds, but also as members of social groups that

have developed their own cultural evolutionary pathway. We use a new methodology

for coding gestural sequences developed by the first author from footage collected by

the second author during two field trips in Grand Terre, New Caledonia. We set out to

establish whether repeated interactions between fledged juvenile birds and

accompanying adults in the context of using tools to forage, can be considered as a

recurring form of learning or even a teaching process. The theory models include

ecological psychology and dynamic cognition. They allow us to consider learning

processes as dynamics of directed, selective, variable and adaptive physiological and

neuronal plastic change which can only occur when the learner is embedded in an

appropriate social, cultural and ecological mileu. Our qualitative results show

consistent gestural patterns embedded in interactions between NCC juveniles and

adults. The patterns suggest that juveniles mature into expert tool-users because they

are cognitively capable of learning. They benefit from having activities graded for them

through time by their adult mentors, and from their own initially unrewarded

perseverance. We end with some brief speculation about the implications of our NCC-

centred findings for evolutionary anthropology.

78

Generalized Darwinism revisited: how a new synthesis changes our view on cultural

evolution

Karim Baraghith

Düsseldorf Center for Logic and Philosophy of Science, Heinrich-Heine University,

Germany

Theories of cultural evolution, dual inheritance theory or generalized Darwinism have

been around for some decades now. The idea is that cultural systems obey the same

abstract principles of variation, selection and reproduction in their development as

biological systems - organisms, genes and populations - do. As well as heavy

enthusiasm, like the ability to provide a long expected scientific synthesis for the social

sciences (resembling the “modern synthesis” for the life sciences, cf. Mesoudi 2011),

such approaches faced severe skepticism and critiques from various directions.

Exceptional aspects of cultural transmission - like conformist bias, blending inheritance

or guided variation - led some researchers doubt that cultural evolution is truly

“evolution”. Furthermore, the identification problem of cultural units or variants

(memes) and of cultural “generations” seemed to undermine the application of

population dynamical or phylogenetic methods for the study of cultural phenomena.

But meanwhile, our biological understanding of "evolution" itself has evolved in various

directions as well. Aspects like plasticity, epigenetic regulation, inclusive inheritance

and niche construction seemingly change our view on evolution in direction of an

“Extended Synthesis” (cf. Pigliucci/Müller 2005). In this paper, I am going to show that

most approaches in the field of cultural evolution today still adopt and assume a very

classical notion of evolution as e.g. given by population genetics. I want to argue that

many critiques are loosing grip, when we involve a more fine grained notion of cultural

“evolution”, which takes into account these new findings and makes evolutionary

models much more applicable to the study of sociocultural systems. I am going to

demonstrate that in three case-studies.

79

Stress-induced mutagenesis systems and the adaptive immunity

Bartlomiej Swiatczak

University of Science and Technology of China, China

Stress response is a basic survival reaction in all living things from bacteria to humans.

In the case of evolving cells, it often takes the form of genome restructuring, which

amplifies variation and accelerates the process of collective adaptation of these cells to

altered environmental conditions. While stress-enhanced evolution is suppressed in

most bodily cells due to their subordination to somatic duties, certain cells of

multicellular organisms like cancer cells and adaptive immune cells do evolve and along

with reclaiming their evolvability, they reemploy their genome restructuring capacities

to diversify their protein products in response to stress or danger. The aim of the

presentation is to show that due to their potential to provide population-level

protection from environmental threats the conserved mutagenesis systems act as

primordial systems of defense for these populations. In fact, stress and danger

responses have been considered key to immunity and unicellular communities are

already able to generate ones. Harnessed by vertebrate organisms, these ancient

danger response systems extend their protective effects from the cell population to the

organism as a whole. Overall, an attempt is made to demonstrate that instead of being

a new invention in the history of immunity, stress-induced hypermutation is its

primordial form, which resuscitated in vertebrate immune systems helps to direct

somatic evolution of adaptive cells towards an adaptive success.

80

Phenotypic integration as both developmental bias and developmental facilitation of

evolutionary diversification: empirical examples from teeth, toes, and torsos

Kathryn Kavanagh

University of Massachusetts Dartmouth, USA

Most theory considering developmental bias on evolution presumes that

developmental integration during morphogenesis acts to reduce variations in a

population, which then biases evolution since selection can only act on variation that

exists. Here I show that it can act as both bias and facilitator of morphological

diversification. An empirical example will be presented of developmental integration

among segments via an activator-inhibitor based mechanism (the Inhibitory Cascade),

which leads to biases in evolution of segment proportions in diverse tissues such as

molars, phalanges, and somites. However, we will also present a study which shows a

counter example, where developmental integration among adjacent tissues (jaw and

tooth) facilitates novel variations and thereby increases evolutionary diversification.

Other examples will show that, in some cases, a large number of rib cartilage variants

are maintained in a population at an extraordinarily high frequency, while, in other

cases, cartilage initiation pattern is extraordinarily stabilized, even when there is

apparent selection against the stable form. This two-sided role of developmental

integration in evolution, and how it affects evolutionary theory, will be explored.

81

Can plasticity create irreversible constraints?

Alfredo Rago, Miguel Brun-Usan, Kostas Kouvaris, Tobias Uller, Richard A Watson


Many traits in nature show no or limited viable variability. This lack of variability

suggests the existence of fundamental boundaries to evolution, known as

developmental constraints. While constraints can result from non-evolvable physical

and chemical properties of life, many biological features that are the result of past

evolution are also highly constrained. This can be explained if evolving in one direction

creates constraints against moving backwards. For example, evolving a new trait may

eliminate the ability to produce viable variants that remove the trait.

While empirical observations overwhelmingly support that evolution can be

irreversible, few theoretical models try to explain how new constraints evolve, and why

they prevent reversible evolution. Here we focus on the hypothesis that developmental

interactions cause constraints. To do so, we simulate the evolution of developmental

plasticity using a developmental model based on gene regulatory networks (GRNs).

We demonstrate that our developmental model can evolve traits that respond to

environmental cues (developmental plasticity) according to simple rules, such as

proportional increase and non-linear dynamics. Evolution of developmental plasticity

changes developmental organization in a way that causes genetic mutations to have

similar phenotypic effects as changes to environmental inputs. Conversely, mutations

with phenotypic effects that are different from plastic responses become rare (bias).

Finally, we show how these biases constrain evolution to proceed in the direction of

plasticity even when alternative directions would provide greater fitness, and prevent

the selection of less biased mutants.

Our work thus highlights how models that integrate general assumptions about

development can explain the evolution of constraints in present-day organisms. This

reduces the need to invoke ad-hoc constraints and helps us understand when

evolutionary changes are expected to be irreversible.

82

Cultural evolution, revolution and transmission of humpback whale song

Ellen C Garland


Culture, the sharing of behaviours or information within a community acquired through

some form of social learning from conspecifics, represents a ‘second inheritance

system’. Male humpback whales (Megaptera novaeangliae) sing a long, stereotyped,

and culturally transmitted song display. At any point in time most males within a

population will sing the same version (arrangement and content) of this complex sexual

display. However, the song is continually evolving and males must constantly learn and

incorporate these changes into their own song to maintain cultural conformity. In

addition to evolutionary change, song also undergoes radical ‘revolutions’ where a

novel song introduced from a neighbouring population rapidly and completely replaces

the existing song. Multiple humpback whale song revolutions have spread across the

South Pacific region from the east coast of Australia across to French Polynesia, with a

one to two year delay. This has occurred regularly, rapidly and repeatedly across the

region; however, we still have a limited understanding of the underlying mechanisms

driving this cultural phenomenon and its implications for extra-genetic inheritance.

Using empirical data, I will present our current understanding of the mechanisms

involved in the song learning process, how these processes may be disrupted, and

finally the evolutionary implications for this cultural phenomenon.

83

Niche construction: resolving the process debate

Tim Lewens


The lively debate going on among evolutionists regarding the nature and significance of

niche-construction is seamlessly scientific, historical and philosophical. Many scientists

also consider the debate’s outcome to have significant implications for biological

practice. In this talk I assess three claims about niche-construction. The first (from some

niche-construction enthusiasts), is that if we are to explain adaptation, we must

supplement natural selection with an additional force that runs from organism to

environment. The second (from some niche-construction sceptics) is that while

selection (and other basic evolutionary processes) have direct impacts on evolution, the

influence of niche-construction is only indirect. The third (from the most prominent

enthusiasts) is that niche-construction is a fundamental process of evolution that has

not been recognised as such, but which has importance for our understanding of

adaptation. I cast doubt on the first two claims. Regarding the third, I draw on Fisher’s

case for runaway sexual selection to point to a pragmatic strategy that can secure niche

construction as an important evolutionary 'process', regardless of whether it is also

judged to be ‘fundamental’.

84

Comprehensive transcriptomics tracks changes in gene expression and coding

sequences during C4 evolution

Lamiaa A Munshi, Pascal-Antoine Christin

The University of Sheffield, UK

C4 photosynthesis is a novel adaptation that increases plant productivity in tropical

conditions. It is a complex trait requiring the coordination of many biochemical,

anatomical and genetics components. Despite its complexity, the C4 trait evolved more

than 60 times independently in flowering plants. However, the history of genetic

modifications leading to the C4 trait remain poorly understood. The Molluginaceae

family includes closely related species that span C4 photosynthesis as well as a variety

of non-C4 types. Capitalizing on this system, we used comparative transcriptomics to

track the changes gene expression levels and coding sequences along the phylogenetic

tree. Genes encoding core C4 enzymes are upregulated in C4 accessions, as expected.

However, we show that these genes were already upregulated in some non-C4

ancestors, which likely facilitated transitions to a C4 biochemistry. By sampling multiple

populations per species, we were further able to demonstrate that amino acid

replacements adapting the proteins for the C4 function followed the upregulation of

genes, and continued when the plants were already C4. Our comparative work

therefore shows how a complex biochemical pathway can be gradually assembled

through multiple rounds of changes in gene expression and coding sequences.

85

Computational complexity is an ultimate constraint on evolution

Artem Kaznatcheev

University of Oxford, UK

Experiments show that evolutionary fitness landscapes can have a rich combinatorial

structure due to epistasis. For some landscapes, this structure can produce a

computational constraint that prevents evolution from finding local fitness optima --

thus overturning the traditional assumption that local fitness peaks can always be

reached quickly. I introduce a distinction between easy landscapes of traditional theory

where local fitness peaks can be found in a moderate number of steps and hard

landscapes where finding local optima requires an infeasible amount of time.

Hard examples exist even among landscapes with no reciprocal sign epistasis; on these

semi-smooth fitness landscapes, strong selection weak mutation dynamics cannot find

the unique peak in polynomial time. A candidate example of such a landscape is yeast’s

tRNA gene for recognizing the arginine codon AGG.

To fully model myriad proximal constraints on adaptation (methods of generating

variation, population structure, stochasticity, etc), I consider all polynomial time

algorithms implementing a step of evolutionary dynamics. On hard rugged fitness

landscapes that include reciprocal sign epistasis, even these arbitrary evolutionary

dynamics cannot find a local peak quickly. This negative result allows reasoning

rigorously from disequilibrium to establish the positive results that – even on static

landscapes – costly learning can remain adaptive and hitchhiking can maintain

cooperation for-effectively-ever.

Moreover, on hard landscapes, the fitness advantage of nearby mutants cannot drop

off exponentially fast but has to follow a power-law that long term evolution

experiments have associated with unbounded growth in fitness. Thus, the constraint of

computational complexity enables open-ended evolution on finite landscapes. Knowing

this constraint allows us to use the tools of theoretical computer science to characterize

the fitness landscapes that we expect to see in nature. Just how ubiquitous hard

landscapes are in nature remains on open empirical question.

86

Population-level variation in sperm-mediated paternal effects in threespined

sticklebacks

Jennifer Hellmann, Alison Bell

University of Illinois Urbana-Champaign, USA

In addition to genetic adaptation and developmental plasticity, parental experiences

can have lasting effects on offspring phenotypes (parental effects). This has long been

appreciated with respect to maternal experiences; however, only recently have we

realized that paternal experiences can be transmitted to offspring through multiple

mechanisms, including via modifications to sperm. There are many outstanding

questions regarding the function and ubiquity of these sperm-mediated paternal

effects. Theory predicts that offspring should attend to parental cues when they are

reliable and informative of future environmental stressors, but should ignore parental

cues when parental environments are not highly correlated with offspring

environments. To understand if there is variation in the extent to which parental cues

are integrated into offspring phenotypes, we assessed the direction and magnitude of

sperm-mediated paternal effects across populations of Alaskan threespined stickleback

(Gasterosteus aculeatus). We found no significant effect of paternal exposure to

predators in a newly established freshwater population of sticklebacks. However, in a

marine and well-established freshwater population, we found that paternal exposure

to predation risk impacted both the variance and mean of offspring traits. For both

populations, variance in behavioral and morphological traits was greater in offspring of

predator-exposed fathers compared to offspring of control fathers. However, the shift

in the mean values of offspring traits differed across the two populations; for example,

offspring of predator-exposed fathers were less active than offspring of control fathers

in a marine population, but in the well-established freshwater population, offspring of

predator-exposed fathers were actually more active than offspring of control fathers.

These data demonstrate that there is population-level variation in both the direction

and magnitude of paternal effects, suggesting that environmental conditions and

genetic cues influence the extent to which parental cues alter offspring phenotypes and

potentially facilitate adaptation to environmental change.

87

The effect of parental care on population evolvability

Sonia Pascoal, Hideyasu Shimadzu, Rebecca M Kilner


A population’s evolutionary history potentially affects its capacity to evolve because

past selection events determine the extent of current genetic variation. Many animals

are exposed to selection from a social environment - the conspecifics with whom they

routinely interact. However, the effect of past selection by the social environment on

current genetic variation is poorly understood. Here we use experiments with the

burying beetle Nicrophorus vespilloides to investigate whether selection by parents

changes the extent of genetic variation in a population. We measured mutation load by

inbreeding individuals from replicate experimental populations that had experienced

different levels of care for 25 generations. We tested two predictions: 1) if parental care

conceals the negative effects of new deleterious mutations, then inbred individuals

with a history of parental care should be more likely to go extinct than those with a

history of no care; 2) if parental care conceals the positive effects of new beneficial

mutations, then inbred individuals with a history of parental care should be more

resilient to inbreeding than those with a history of no care. Our results support the

former prediction. We conclude that parental care influences the quality of a

population’s standing genetic variation, so changing the raw material available for

future evolutionary adaptation.

88

The thrifty watchmaker

Michael Lachmann, Wim Hordijk

Santa Fe Institute, USA

Why do we need to exercise to keep our muscles in shape? Why not develop the right

muscles without such input? Wolff’s law states that bones adapt to the stresses placed

on them. A similar law, Davis’s law applies to soft tissue. Plants use light to direct their

growth, and blood vessels oxygen concentration. In these cases the underlying

developmental mechanism seems to “understand” its high level function. How did

evolution reach that fitness peak, when it can only see the local benefits to the lineage?

In my talk I will claim that a key to understanding all these and other related phenomena

is to switch our view of what a genotype is. Instead of thinking of a genotype as coding

for a phenotype, or maybe a G by E matrix coding for a phenotype, which should think

of the genotype as an interpreter of environmental information into phenotypic

information. The genotype is interpreting the micro-environments that the

developmental process encounters to form the phenotype. This view allows us to

understand why developmental processes seem so general.

89

Variable phenotypic plasticity and its consequences for evolutionary potential

Franziska S Brunner, Alan Reynolds, Ian Wilson, Stephen Price, Steve Paterson, David

Atkinson, Stewart J Plaistow

University of Liverpool, UK

It is often hypothesized that the pattern of phenotypic plasticity in a population may

direct the course of subsequent evolution. As a result, plasticity in populations is often

summarized as an average response and labelled as being either adaptive or

maladaptive. However, this approach ignores the genetic variation in phenotypic

plasticity present within populations and assumes that the mean plastic response is

representative of the whole population. Whether this is acceptable or not depends on

both the magnitude and the type of plasticity variation maintained in populations;

something that is rarely measured.

In this study, we quantified the multivariate life history plasticity of 43 clones collected

from the same population in response to both temperature and resource level

differences. We calculated population-level variation in both the phenotypic direction

and magnitude of plastic responses and compared this to two reference populations

from very different latitudes. In all cases, we found that within population variation in

plastic responses is high and context-dependent. Based on these findings, we discuss

the reasons for the maintenance of plasticity variation within a population and its

consequences for evolutionary potential. We conclude that there may be many

instances where the effect of phenotypic plasticity on the rate and direction of

evolution is not accurately represented by an average reaction norm.

90

Comparative genomics of convergent evolution in the emergence of complex traits

Claudio L Flores Martinez

University of Hamburg, Germany

With the ever-increasing number of sequenced metazoan genomes, it becomes

apparent that several complex traits, which had long been held to have evolved in a

homologous manner, emerged convergently in distantly related animal lineages.

Complex body plans, nervous systems and eusociality certainly are among the most

interesting traits that – under the scrutiny of comparative genomics and functional

analysis – are breaking away from the hegemony of homology. However, the genomic

basis of convergent evolution is not well characterized. Nevertheless, it constitutes a

decisive element in understanding the prevalence and adaptive value of convergence

throughout metazoan bio-complexity. This first systematic and theoretical study on the

genomic underpinnings of convergent evolution focuses on the current rewriting of

nervous system evolution in Metazoa. More generally, at least four major mechanisms

driving convergent evolution of metazoan genomes can be distinguished: 1) protein

family expansion, 2) convergent amino acid substitution, 3) de novo recruitment and 4)

transposable elements. Information and (biological) systems theory are reframing the

phenomenon of convergent evolution in terms of top-down causation (the process

whereby higher levels of organization in structural hierarchies constrain the dynamics

of lower levels) and inferred functional equivalence classes. This project explores the

convergent emergence of complex traits across multiple scales and at different levels

of organizational bio-complexity (from molecular instantiation to cell type, tissue and

beyond) within a phylogenetically-inspired context. It aims at establishing a coherent

approach allowing, first, to identify problems in evolutionary biology that can be solved

by using comparative genomics; second, outline the kind of bioinformatic pipelines

needed for solving the problem (e.g. protein family expansion vs. amino acid

substitution) and, third, interpret these results within newly forged theoretical and

quantitative frameworks that deal with the highly dynamic genomic processes and

mechanisms underlying the convergent emergence of complex traits.

91

Epigenetic potential in native and introduced populations of house sparrows (Passer

domesticus)

Haley E Hanson, Holly J Kilvitis, Aaron W Schrey, J Dylan Maddox, Lynn B Martin

University of South Florida, USA

Epigenetic potential, defined as the capacity for epigenetically-mediated phenotypic

plasticity, may play an important role during range expansions. During range

expansions, populations may encounter relatively novel challenges, including novel

pathogens, while experiencing lower genetic diversity due to bottlenecks and/or

founder effects. Phenotypic plasticity may allow individuals to rapidly cope with these

challenges. Here, we asked whether one form of epigenetic potential (i.e. the

abundance and/or position of CpG sites within gene promoters) varied among native

and introduced populations of house sparrows Passer domesticus in three microbial

surveillance genes: Toll-like Receptors 1B (TLR1B), 2A (TLR2A), and 4 (TLR4). Evidence

suggests that increased expression of TLR2 and TLR4 may play a role in house sparrow

range expansions, and that methylation at one CpG site within the putative promoter

region of TLR4 was a strong predictor of TLR4 expression in house sparrows. We

characterized i) total CpG sites, ii) total gains of CpG sites, and iii) losses of CpG sites in

TLRs 1B, 2A, and 4. We hypothesized that introduced populations would have more

total CpG sites, more gains of CpG sites, and fewer losses of CpG sites (or overall higher

epigenetic potential). We found that introduced populations had a higher total number

of CpG sites in TLR2A and TLR4, but not in TLR1B. Introduced populations had more

losses of CpG sites in TLR1B, more gains of CpG sites in TLR2A, and fewer losses of CpG

sites in TLR4 compared to native populations. These trends were not driven by genetic

differences between populations, nor by genetic diversity. This suggests that selection

is acting on CpG sites in introduced populations, but possibly through different

mechanisms, and that epigenetic potential may be playing a role in house sparrow

range expansions.

92

Why does natural selection favour the transgenerational inheritance of social

adversity?

Bram Kuijper, Rufus A Johnstone

University of Exeter, UK

Numerous studies have shown that social adversity in early life can have long-lasting

negative consequences for social behaviour in adulthood, consequences that are often

propagated to future generations. Given these transgenerational effects, it is puzzling

why natural selection might favour such sensitivity to an individual's early social

environment. To address this question, we model the evolution of social sensitivity in

the development of helping behaviours, showing that natural selection indeed favours

individuals whose tendency to help others is dependent on early-life social experience.

We find that natural selection typically favours positive social feedbacks, in which

individuals who received more help in early life are also more likely to help others in

adulthood, while individuals who received no early-life help develop low tendencies to

helping others later in life. This positive social sensitivity is favoured because of a

transgenerational relatedness feedback: patches with many helpers tend to be more

productive, leading to higher relatedness within the local group, which in turn favours

higher levels of help in the next generation. By contrast, patches with few helpers tend

to be less productive, leading to lower relatedness, which in turn favors lower levels of

help in the next generation.

93

Trait mean-variance relationships impact speed of adaptation

Rebecca B Hoyle, Thomas HG Ezard


Quantitative genetics models of trait evolution typically assume that trait (co)variances

remain constant owing to mutation-selection balance. Mean-variance relationships are,

however, common in biological systems. Morphological trait data from fossil planktonic

forminifera suggest that the trait variance may vary linearly with its mean. We present

a quantitative genetics model of trait evolution in which the trait variance changes

adiabatically with the mean and show that this leads to faster trait evolution and long-

term maladaptation following an environmental shift. This raises interesting questions

about the extent to which evolutionary trajectories favour speed of adaptation to

changing environmental conditions over fitness outcomes and the limitations of the

ubiquitous assumption of constant trait variances through mutation-selection balance.

The influence of multi-level demographic and ecological feedbacks in the evolution of

cooperation

Antonio MM Rodrigues


Theoretical models of social evolution typically seek to uncover which genetic and

ecological factors mediate the evolution of cooperative behaviours. In addition, most

models assume that these behaviours have small phenotypic effects and therefore the

interdependency between the behaviour and the environment is considered to be

minimal. More generally, behaviours will lead to significant changes in phenotypes,

which have the potential to impact both the local and the global environment, which,

in turn, will feedback to change the evolutionary trajectories of cooperation. These

processes, which take place at different levels of biological organisation, are still poorly

understood. Here, I will outline a model that captures the effect of cooperative

behaviours at different levels, from the individual to the population, and therefore

includes the impact of ecological feedbacks on social phenotypes. I will show how the

evolution of cooperative behaviours modifies local demography as well as population

density across multiple-generations and I will describe how these multi-level processes

can influence the evolutionary trajectories of behaviours. Moreover, I will analyse cases

in which individual conditions can be transmitted over multiple generations, and I will

show how this can either prevent or promote the evolution of cooperation.

94

The evolution of teaching and anti-teaching

James Allen, Rufus A Johnstone


Alongside genes, culture is also transmitted between generations. Whilst both empirical

and theoretical work has established social learning in animals as crucial for the

transmission of skill, definitive examples of teaching are rare, and what examples there

are exist in very different species. Defining teaching as an enhancement of social

learning, we develop a theoretical framework of skilled individuals in a structured

population, and investigate two outstanding issues within the evolution of teaching: the

importance of demographic competition, and the prevalence of cooperative breeders

within empirical examples.

By including the cost of demographic competition, we pinpoint the existence of a novel

adaptive behaviour: the costly reduction of other’s social learning. In parameter

regimes where competition dominates (high vertical transmission and low philopatry)

a reduction of social learning is selected for, a behaviour we label anti-teaching. Just as

is observed for teaching, we find that anti-teaching is most adaptive when the skill is

highly adaptive and difficult to learn. Introducing a trade-off between teaching effort

and mortality, anti-teaching is adaptive even when it is costly to reduce the learning of

others. Notably, when competitive considerations dominate, individuals are willing to

reduce the social learning of others even at a cost to their own mortality.

Furthermore, we extend our framework through the inclusion of age-dependent

teaching efforts, enabling us to study the age at which teaching is adaptive, and the

effect of reproductive skew on the evolution of teaching. By introducing the novel

behaviour described above, and age-specific teaching efforts, this work contributes to

our understanding of the evolution of teaching, and more generally the spread of

culture in animals.

95

Evolution of small RNA production under fluctuating environmental conditions

Willian T.A.F. Silva, Sarah P Otto, Simone Immler

Uppsala University, Sweden

Recent empirical studies have addressed the importance of small RNAs (sRNA) not only

in their primary function of post-transcriptional regulation of gene expression but also

in their ability to respond to fluctuations in environmental conditions and be

transmitted across individual generations, creating a mechanism of non-genetic

inheritance. Small RNA function has been particularly associated with an organismal

response to suboptimal environmental conditions caused by biotic (e.g. viral infection)

or abiotic factors (e.g temperature fluctuations) and its transgenerational, potentially

adaptive effect. We created a model of sRNA dynamics including the major biological

components involved in sRNA biosynthesis via the ping-pong amplification process that

is characteristic of Piwi-interacting sRNAs. Additionally, we include a transgenerational

effect and responsiveness in amplification rates, and explore the sRNA dynamics across

individual generations under different fluctuating environmental conditions. Our

results show that mechanisms of sRNA amplification become highly invasive when

coupled with responsiveness to environmental conditions, and responsiveness in the

soma or in the germline can result in different evolutionary outcomes depending on

levels of environmental correlation. We consider our findings in the context of current

empirical studies and discuss the possible evolutionary consequences of non-genetic

inheritance.

96

Inheritance of acquired traits - implications for evolution?

Sophie Juliane Veigl

University of Vienna, Austria

The inheritance of acquired traits (IAT) has regained prominence in the current heyday

of epigenetics. The transmission of acquired information from parents to offspring,

even for several generations has been reported in a wide range of species. Proponents

of the Extended Evolutionary Synthesis (EES) argue that modes of inheritance and

evolution, which are not Neo-Darwinian, are certainly possible. Defenders of the

Modern Synthesis (MS) mobilize several arguments to counter such claims. For

instance, they argue that IAT has no bearing on evolutionary timescales, as it is too

short-lived! Or, they argue that no matter what the specific mechanism of IAT is, all

effector molecules are encoded in DNA, and thus subjected to random mutation and

natural selection. Is IAT as a mode of evolution thus even plausible? What can be said

in its defence?

In this paper I discuss two scenarios in which IAT might have an impact on evolution. I

use the example of the inheritance of RNAi in the nematode C elegans. Specifically, I

present two arguments that address criticisms of defenders of the MS. First, I point to

amplification systems in C. elegans, that prolong IAT for up to 80 generations, and

counting. Thus, evolutionary timescales of IAT are possible, at least in principle. Second,

I consider the active role of small RNAs on restructuring the genome: small RNAs alter

the susceptibility of individual stretches of DNA for random mutations, by making them

more or less accessible. Thus, they modulate the impact of classical modes of evolution.

Based on these two examples I conclude that by looking at the mechanisms underlying

IAT, arguments of defenders of the MS can be rejected. Focusing on the specific

mechanisms also provides incentives for testing the role of IAT in evolution.

97

Egg development plasticity and the transmission of maternal effects in Daphnia

Stewart Plaistow, Aishah Aslam, Megan Hasoon, Marco Marcello

University of Liverpool, UK

Environment-induced non-genetic inheritance (NGI) may play a key role in facilitating

rapid adaptation to changing environments. However, the mechanisms underpinning

NGI remain obscure and we currently know little about how genetic, non-genetic and

environmental cues are integrated. The egg phenotype is critical in this respect because

it provides many routes for information transfer from one generation to the next and

defines the initial environment that inherited genes are exposed to. Parthenogenetic

organisms such as Daphnia are ideal models for investigating the integration of

developmental cues because large numbers of genetically identical individuals can be

reared in different environments in parental and offspring generations, making it easy

to separate genetic and non-genetic influences. However, variation in egg phenotypes

is typically only ever studied at a simplistic level. Here we use metabolomics to compare

the egg phenotypes produced by mothers exposed to different environments, or

mothers occupying different states (sexual vs asexual, old vs young). We then use

confocal microscopy to compare the embryology of offspring from mothers from

different environments, or different states. Our results suggest that (1) changes in the

maternal environment or state result in changes in the egg phenotype, and (2) that

changes in the egg phenotype results in egg development plasticity that can explain life-

history variation in adults. We conclude that offspring provisioning is likely to be a key

factor explaining how genetic, non-genetic and environmental cues are integrated.

98

Environmental heterogeneity mediates the evolution of trans-generational effects on

fitness

Martin I Lind, Martyna K Zwoinska, Johan Andersson, Hanne Carlsson, Therese Krieg,

Tuuli Larva, Alexei A Maklakov


Theory maintains that environmental heterogeneity can shape the evolution of trans-

generational parental effects on offspring fitness but the experimental evidence is

limited. Constant or predictably fluctuating environments should favour the evolution

of adaptive trans-generational parental effects. In contrast, the absence of correlation

between parental and offspring environments should select against trans-generational

effects. Here we investigated the role of environmental heterogeneity in the evolution

of trans-generational parental effects using experimental evolution in response to

temperature. Genetically diverse populations of the outcrossing nematode worm

Caenorhabditis remanei, adapted to 20°C, were exposed to a novel, stressful high

temperature (25°C) for 30 generations, in constant, increasing or fluctuating

temperature regimes, with either strong or zero correlation between parent and

offspring temperature environment.

We found evolution of trans-generational effects, since fitness in the novel, high

temperature depended upon parental environment. While lines evolving in correlated

environments required parental exposure to high temperature to achieve high

reproduction in this environment, the total reproduction of lines evolving in

unpredictable environment was independent of parental environment. This was caused

by the adaptive maintenance of an anticipatory trans-generational parental effect in

lines evolving in correlated environments, while this trans-generational inheritance had

been adaptively lost in lines evolving under non-correlated environmental

heterogeneity. Moreover, the individual fitness (lambda) of the lines evolving in

uncorrelated environments was actually highest when parents and offspring where

grown in opposite environments (simulating environmental fluctuations). Therefore,

environmental heterogeneity influences the evolution of trans-generational

inheritance.

99

Abstracts: Poster Presentations

P01

Expanding the conceptual scope of niche construction theory

Bendik Hellem Aaby, Grant Ramsey

KU Leuven, Belgium

Niche construction, commonly defined as organisms actively changing selection

pressures by altering the feature-factor relationship between themselves and their

environment, has become increasingly incorporated into evolutionary dynamics in

contemporary evolutionary biology. The interest in niche construction was largely

sparked by Lewontin’s work on biological constructivism, and more recently builds on

niche construction theory (NCT) Odling-Smee et al. (2003). NCT argues that we can

distinguish two fundamental kinds of niche construction—perturbative and

relocational. These concepts refer to the way in which niche construction is brought

about by the activities of the organisms. ‘Perturbative’ captures the cases in which an

organism’s activities alter its physical (external) environment and thereby changes the

selection pressures, e.g., ecological engineering. ‘Relocational’ captures the cases

where the organisms relocate in space (without modifying the environment), and

thereby changes the selection pressures, e.g., migration.

The NCT conception of niche construction is considerably narrower than that of

Lewontin. In this talk we defend a broader framework more in line with Lewontin’s

original account. We hold that there are three fundamental ways in which organisms

can construct their niche: ‘constitutive’, ‘relational’, and ‘external’ niche construction.

We aim to show that NCT’s ‘perturbative’ and ‘relocational’ excludes important niche

construction cases, and we suggest that our framework is able to subsume all the

phenomena described by NCT while also allowing us to account for far more cases of

niche construction, especially social niche construction. We end our talk by showing

how this broader Lewontian framework can help us elucidate how niche construction

can be treated as an evolutionary process as understood by the extended synthesis.

100

P02

Could a major evolutionary transition have happened in human evolution?

Paulo C Abrantes

University of Brasília, Brazil

Paulo C. Abrantes attempted to make explicit, in a 2013 paper, the conditions under

which a major evolutionary transition could have taken place in human evolution by

using Godfrey-Smith's criteria for telling marginal from paradigmatic Darwinian

populations in the cultural domain. In that paper, I argue that the emergence of

cooperation in hunter-gatherer bands during the Pleistocene could have been a major

transition in individuality, associated with the evolution of a species-specific social

psychology.

Difficulties arise, however, when one attempts, by adopting this approach, to come up

with a notion of fitness pertaining to cultural groups, among other problems in applying

to those groups the requirements for biological individuality. I will examine in this talk

the plausibility of another scenario, in which a major transition in individuality could

have happened later, associated with ultrasociality in large-scale societies. Godfrey-

Smith's conceptual scheme applies less problematically in this case. This scenario

doesn't presuppose any further psychological capacities on top of those which emerged

among hunther-gatherers, arguably by gene-culture coevolution. A major transition

could have been the outcome of just cumulative cultural evolution. Furthermore, this

scenario has the advantage to be compatible with other attempts to reconstruct the

latest stages of the evolution in the Homo genus, including those proposed by Maynard

Smith and Szathmáry, Turchin, Richerson, Boyd, and Tomasello, among others.

However, this alternative scenario has to deal with criticism from those that reject

adaptationism and push for an expanded version of the theory of evolution in able to

integrate developmental and evolutionary processes. A more heterodox proposal along

those lines would attempt to explain human ultrasociality by underlining the role of

ontogenetic processes and niche construction, and playing down natural selection in

accounting for that trait.

101

P03

Dialectics of evolution; epistemology versus practice

Ozan A Altinok

WWU Münster, Germany

My main aim is to distinguish the usages of evolution such as evolutionary models,

evolutionary theory, evolutionary phenomena and “evolution proper” to understand

the extent of change in the call for change in the Modern Synthesis (MS) in favor for the

Extended Evolutionary Synthesis (EES). Gould (1981) distinguishes evolution as fact and

theory. I would like to add evolution as instrument to have a broader perspective to

understand change in evolution. Through elaborating that scientists often make the

distinction between the theories which are “home truths” (Hacking, 1984) and theories

which are under investigation, I will analyze the limits and the location of change in

“evolution” that is happening and that is expected to happen by EES.

Wray et. al, (2014) call for an understanding of evolution which permits the domains to

be on its core, as long as they are empirically validated, while on the very same

commentary Laland et. al (2014) call for an EES for biologists to grasp a better

understanding of evolution. I would like to consider scientific practice more central

rather than theories. Going away from “high theories”, Cartwright (1983), helps us to

see the extent of flexibility in the usage of the concept of evolution, holding that

evolutionary theory, as a theory, has limited relevance to the practices of biologists. I

claim that scientists are generally very pragmatic and instrumentalist (Matthews, 2017),

when considering their work in most cases, as guidelines of research.

I claim that, Callebaut’s (2010) understanding of dialectics of unity and disunity should

be extended to dialectics of practice of biology and biological theory with the scientists

in the center to grasp the possibility of change in accurate aspects of evolution through

EES and other possible scenarios that are in the form of a movement, not only a protest.

102

P04

Overprinting and the evolvability of novel protein-coding genes

Zachary Ardern, Stefan Wichmann, Siegfried Scherer

Technical University of Munich, Germany

As techniques to discover short proteins have improved, many overlapping genes

encoding protein products have been discovered in viral, prokaryotic, and eukaryotic

genomes. Methods such as ribosome profiling, large-scale analyses of sequence

constraint made possible by modern databases, and improvements in proteomics have

all contributed to this substantial increase in our ability to detect protein products.

Studying the origin of these genes provides an important case study in evolutionary

innovation. This class of genes is particularly useful for obtaining insights on gene origin

as the genomic context prior to an overprinting event can be inferred with more

confidence than is possible in the case of most intergenic sequences. Important factors

in facilitating the origin through overprinting and subsequent conservation of novel

protein coding genes include contributions of the pre-existing ‘mother’ gene’s

sequence, the reading frame of the overlapping ORF, and the structure of the standard

genetic code. Through a combination of phylostratigraphy, ancestral sequence

reconstruction, and modelling of overlapping gene construction and alternative codes,

we report on the contributions of each of these factors, placing them within a broader

context of genomes’ inherent potential for evolutionary innovation.

103

P05

Morphological divergence in Drosophila ananassae due to nutritional variations

through changes in allometric relationship and trait size

Bhumika, Arvind Kumar Singh

Banaras Hindu University, India

Developmental plasticity in response to nutritional changes plays a substantial role in

adaptive evolution and diversification of morphological traits in organisms. We

investigated the diversification in wing dimensions and its relative scaling with respect

to body size, i.e., allometric scaling in response to dietary variations in carbohydrates

and proteins quantity in Drosophila ananassae. For the experiments, ten isofemale lines

of D. ananassae derived from Bilaspur, India and karyotypically characterized stocks of

D. ananassae from Varanasi, India were utilized. We found that protein and

carbohydrate variations in dietary media induced significant genetic variation in

plasticity for allometric slopes of wing traits in males and females of Bilaspur

population. However, Varanasi population refrained from showing such effect in

response to dietary variations. The results suggest that changes in allometric

coefficients due to dietary variation is population specific and this may lead to

difference in morphological divergence in populations, thereby, causing evolutionary

allometry. The analysis of reaction norm graphs suggested the involvement of alpha

inversion present in 2L chromosome of Varanasi population, in conferring adaptation

to different nutritional environmental conditions due to changes in static allometry of

different wing dimensions. We detected that effect of dietary protein variations on

plasticity of absolute body size and wing dimensions was highly genotype dependent

when compared to variation in sucrose quantity, in both the sexes and in both the

populations. The results signify that variations in protein in natural environment have

greater tendency to induce selective pressure on absolute trait values of genetic

variants as compared to carbohydrates and thereby had greater capability of inducing

morphological diversifications through the process of genetic accommodation. Overall,

this study reveals that the nutritional ecology governs the morphological divergence of

traits size and shape in a genotype dependent manner in D. ananassae thereby resulting

in morphological evolutions.

104

P06

Coral-built structural complexity enhances coral recruitment: niche construction in

action?

Viviana Brambilla, Andrew H Baird, Miguel Barbosa, Inga Dehnert, Joshua S Madin, Clare

Peddie, Maria A Dornelas


Coral larval recruitment is a key process for coral reef persistence. Micro-habitat scale

rugosity enhances settlement success; however, the effect of structural complexity of

the habitat forming coral colonies remains undetermined. Here, we test the effect of

habitat scale structural complexity on settlement and juvenile presence on artificial

tiles. Structural complexity had a positive effect on settlement (+ 15.71 % of settler

presence probability along the gradient), but no effect on juvenile abundance was

detected. While complexity might be trivial for early stages survivorship, our results

suggest that structural complexity as result of coral morphology creates conditions that

facilitate coral settlement and serve as foundation for further studies of coral niche

construction.

105

P07

When are more experiments necessary? Using evolutionary considerations in

investigating animal minds

Simon Brown

Columbia University, USA

In the study of animal minds, available neuroscientific and behavioural data often

underdetermine which of several radically different models of a particular species' mind

is best. Morgan's Canon traditionally offered one way of responding to this challenge,

but it has well-known problems. One alternative, considered by Sober (2005, 2015), is

cladistic parsimony. But as Sober points out, this principle too will often only very

weakly constrain our attribution of mental traits to animals. One possibility here (Sober

2005, Fitzpatrick 2008) is that we should abandon the search for principles of this kind

and treat these issues entirely on a case by case basis. This paper, however, shows that

we can usefully introduce more general principles for combining evidence about pairs

of species, taking account their phylogenetic relations on the one hand and similarities

and differences in their niches on the other. We can thereby systematically determine

how confident we should be in attributing any given mental state to a given species.

The key to doing this will be to understand a general notion of evolutionary dependence

- roughly, considerations about the order in which various traits must appear in

evolution. The generality and utility of this sort of notion has been underappreciated

in the evolutionary sciences of animal minds. With a computational account of a

cognitive trait, we can understand what it might evolutionarily depend on. This notion

can include the numerous cases where a trait of interest might be an exaptation based

on a computationally similar mechanism used for a rather different function. And this

in turn means that we can use evidence from one species to generate predictions about

related species with much more robustness than might be expected, and without the

need for extensive, expensive and invasive experiments for each species of interest.

106

P08

The evolution of unicellularity in a multicellular world

Charlie Cornwallis, Maria Svensson-Coelho, Karin Rengerfors, Lars-Anders Hansson


Explaining how life evolved from simple replicators in a primordial soup to the array of

biodiversity observed today remains extremely challenging for evolutionary biology.

Increases in the complexity of life have occurred through a series of transitions where

solitary replicators, such as genes, cells and animals (simple life forms), committed to

living in groups, such as genomes, multicellular bodies and societies (complex life

forms). However, not all organisms are complex so what explains when transitions

occur? This question has been addressed by comparing complex and simple organisms.

This allows inferences about the changes that take place during transitions, but reveals

little about the adaptations in simple organisms that facilitate and hinder the evolution

of complexity. Uncovering such ‘precursors’ to complex life is extremely challenging, as

it requires identifying the mechanisms underlying complexity and tracing their origins

back over time. Here I will present on-going work that integrates experimental

evolution and comparative work to understand whether there are ‘precursors’ to both

multicellular and unicellular life using the model system of green algae collected from

non-model wild populations.

107

P09

Enactive evolution

Amanda Corris


According to the enactive approach to biological organization, an organism can be said

to enact or ‘bring forth a world’ through structural coupling, where constructive

interactions between organisms and their environments result in a relationship of co-

evolution between those organisms and their environments as niches. While the

enactive approach mainly aims to provide a theory of cognition, it also provides a

theoretical framework for an alternative to received conceptualizations of adaptation.

I argue that this framework is complementary to niche construction theory and can

bolster a niche construction theory of evolution by supplying conceptual tools that help

to explain the processes involved in the generation of selection. The enactive concept

of evolution by natural drift (Varela, Thompson and Rosch 1991), whereby self-

organizing, structurally coupled configurations of organisms and their niches generate

selection, provides an account of feedback in evolution that can build on similar

accounts offered by niche construction theory. The enactive approach has significant

contributions to make to theories that aim to reconceptualize evolutionary processes

and is worthy of consideration in contemporary debates regarding development and

evolution.

108

P10

Behemoth: Henri Bergson's and Gilles Deleuze's vitalism, a proto evolutionary biology

Jack Robert Edmunds-Coopey

University of Durham, UK

Bergson's philosophy of life and its organicism inspired Deleuze's vitalistic tendencies in

the 20th century which attempt to not only capture life as it is but to use scientific

explanations of life integrated with philosophical ones. The study of life itself is what

fascinated these thinkers which influenced not only their understanding of time in how

the life experience of time provides more insight than a metaphysical account deriving

from Kant. In light in how science despite the instrumental critiques of reason raised

against it by the Frankfurt School in how 'science does not think' in Heidegger's terms

continues to innovate and destroy life as we know it as and not as an object of science.

Thus, it is proposed that the science of life as such is argued to study inanimate things

but if we question like Bergson and Deleuze did this assumption of the binary between

animate and inanimate objects through vitalistic tendencies, arguing that in fact life is

not limited to biological entities but all forms of objects around us in a new

understanding of what constitutes life as such outside a Darwinian and Newtonian

definition which is perhaps restricted to the plant and animal kingdoms. Thus, life as we

know it becomes dissolved if we re-consider the assumptions which delimit animate

things from inanimate things and consider objects as examples of manifestations of life

and how our desires, values, morality and meaning give 'life' to all sorts of objects and

things around us consequently. In conclusion, this paper shall explore the vitalism of

Henri Bergson and Gilles Deleuze and argue that their interactions with reconceiving of

life not only revolutionizes understandings of biology, religion and the human meanings

associated with societies, but that the boundaries and barriers between the humanities

and sciences should not be existent.

109

P11

Effects of phenotypic robustness on adaption and evolvability

Emanuele Rigato, Giuseppe Fusco

University of Padova, Italy

Theoretical and experimental studies have provided evidence for a positive role of

phenotype resistance to genetic mutation, or “phenotype mutational robustness”, in

enhancing adaptation and fostering evolvability. With the aim of contributing to an

understanding of the origin and evolution of phenotypic robustness in living systems,

we adopted a theoretical approach, elaborating on a standard model of evolutionary

dynamics, complemented by computer simulations. Results show that, under common

selective regimes, a certain level of phenotypic robustness is a necessary condition for

adaptation to occur, even in the absence of any drift effects. We also tested the

predictions of the model by exploring some selected features of genome evolution,

through a phylogenetic comparative approach on a sample of 210 eukaryote taxa.

Phenotypic robustness qualifies as major quantitative determinant of biological

system’s evolvability, a key feature of the genotype-phenotype map which would

deserve to be formally included into a more inclusive explanatory framework of the

evolutionary theory.

110

P12

Spadefoot toad tadpoles incur maintenance costs of developmental plasticity in the

form of increased oxidative stress

Pablo Burraco, Miguel A Rendón, Carmen Díaz-Paniagua, Ivan Gomez-Mestre

Doñana Biological Station, CSIC, Spain

Adaptive plasticity enables organisms to improve the match between their phenotype

and the environment under fluctuating environmental conditions. However, despite

this advantage of plastic genotypes, we often observe substantial variation in the

degree of plasticity across genotypes within populations. Costs of maintaining the

sensory and response machinery of plastic traits have been proposed as a plausible

explanation for the maintenance of genetic variation for plasticity, although only a few

studies have detected them empirically. The majority of amphibians have an aquatic

larval phase that typically shows a high degree of developmental plasticity, as they can

alter their growth rate, morphology and behavior against various environmental inputs.

We have tested for physiological maintenance costs of developmental plasticity in larval

spadefoot toads in response to predation risk. We determined the plastic responses of

20 sibships to water-borne predator cues, and estimated parameters associated with

metabolic rate, oxidative stress and immune status on tadpoles under control

conditions. We found that plasticity in response to predator presence entailed

maintenance costs in terms of antioxidant and immune imbalance. Our results indicate

that being plastic can be physiologically costly, suggesting that the role of maintenance

costs in the evolution of plasticity should not be dismissed.

111

P13

The evolution of barriers to exploitative behaviors

Jonathan R Goodman, Paul Ewald


This presentation aims to show how barrier theory, which was generated to understand

how cancer develops through the abrogation of mechanisms that block cellular

reproduction and survival, can be more broadly applied to other systems, including

animal populations and human societies. Barrier theory suggests how Red Queen–like

evolutionary relationships may be ended, or at least paused. Barriers may be

understood as naturally evolved or artificially implemented mechanisms for blocking

the risk of exploitation. Selection will therefore favor those entities capable of

abrogating these barriers: oncogenic viruses, for example, evolved enhanced survival

by blocking the suicide mechanisms of their host cells. We propose a model of barrier

theory suggesting that, for any system, barriers to exploitation and the degree of

exploitation are inversely and logarithmically related. The mode of exploitation may,

furthermore, be unpredicted, which cannot be accounted for in a standard signaling

game model. Modes of exploitation differ depending on the system: invasive organisms

may overcome the evolved barrier to cellular survival and reproduction; a plant may

temporarily escape herbivory by evolving a chemical that none of its herbivores can

detoxify. Together these premises highlight some important consequences for

competitive co-evolutionary relationships.

112

P14

Three lessons of Bergson's creative evolution in the wake of transhuman evolution

Daina Habdankaitė

Vilnius University, Lithuania, and Turin University, Italy

The fast-paced advancement of AI, robotics, gene editing, and other contemporary

technologies is considered by several theorists as the wake of transhuman evolution.

The idea behind it is that human evolution is about to transcend the biological

boundaries of the human organism in order to complete itself in a technically mediated

form. The idea of transhuman evolution brings into question the process of evolution

as well as the notions of species, intelligence, continuity and predictability of the

process of development. By exploring a few examples of AI, robotics, and gene

manipulation in creative practices, I will suggest that Bergson‘s concept of creative

evolution could provide us several valuable lessons in conceptualizing and evaluating

the idea of transhuman evolution.

1) Bergson’s strict division between intellect and intuition, grounded on their relation

to the matter, helps to shine a light on the importance of materiality in the case of

technological advancement of intelligence. To put in technological terms, once

admitted that hardware decides what software is to be developed, is it possible for the

software to influence hardware as well? And if so, what implications does this have on

the conception of evolving intelligence and its transformations?

2) The line that Bergson draws between creativity and calculation helps to crystallize

the problematic aspect of AI development: in order to be considered as AI, it must bear

some resemblance to the human mind; but is creativity something that cannot be

reproduced in and by a machine? What is it like for a machine to be creative?

3) Bergson’s notion of virtuality, when applied in the context of transhuman evolution,

shines some light on the problem of uncertainty that part of AI and robotic technologies

are aimed at solving. Is uncertainty a human fault or an evolutionary system’s default?

113

P15

Evolution of phenotypic complexity: a developmental perspective

Pascal F Hagolani, Roland Zimm, Isaac Salazar Ciudad


How does phenotypic complexity evolve? Indirect empirical evidence suggests that

mutations which decrease complexity are more common than mutations that increase

complexity. Additionally, it has been shown that complex phenotypes have a more

complex relationship between genetic and phenotypic variation, or genotype-

phenotype map, than simpler ones. The view from classical evolutionary biology is that

complex genotype-phenotype maps hinder adaptation. Thus, these complex genotype-

phenotype maps should be rare or, at least, become simpler over the course of

evolution. In this paper, we show that, instead, such complex maps and the above

mutational asymmetry are an inevitable consequence of how genes need to be wired

into networks during development in order to lead to complex morphology.

To explore this question, we randomly wired a huge number of genes and cell

behaviours into networks in EmbryoMaker. EmbryoMaker is a mathematical model

simulating collective changes in cell 3D spatial positions and gene expression due to

such networks. In this way we obtained a “zoo” or an ensemble of in silico 3D embryos.

Real gene networks are not random, but random networks allow an unbiased view on

the requirements for the development of complex phenotypes. We found that the

networks leading to complex phenotypes are: 1) rare; 2) need to be finely tuned to

produce complex morphologies; 3) less stable to noise than networks leading to simple

phenotypes; 4) more likely to lead to a decrease in complexity than to an increase when

they are mutated and; 5) they exhibit a complex genotype-phenotype map. Our results

suggest that, when complexity increases during evolution, it does so at a progressively

slower rate as mutations which increase complexity become rarer, morphologies

become less robust, and a more complex genotype-phenotype map arises.

114

P16

Timing of maternal exposure to toxic cyanobacteria and offspring fitness in Daphnia

magna: implications for the evolution of anticipatory maternal effects

Reinder Radersma, Alexander Hegg, Daniel WA Noble, Tobias Uller


Organisms that regularly encounter stressful environments are expected to use cues to

develop an appropriate phenotype. Water fleas (Daphnia spp.) are exposed to toxic

cyanobacteria during seasonal algal blooms, which reduce growth and reproductive

investment. Because generation time is typically shorter than the exposure to

cyanobacteria, maternal effects provide information about the local conditions

subsequent generations will experience. Here, we evaluate if maternal effects in

response to microcystin, a toxin produced by cyanobacteria, represent an inheritance

system evolved to transmit information in Daphnia magna. We exposed mothers as

juveniles and/or as adults, and tested the offspring’s fitness in toxic and non-toxic

environments. Maternal exposure until reproduction reduced offspring fitness, both in

the presence and in the absence of toxic cyanobacteria. However, this effect was

accompanied by a small positive fitness effect, relative to offspring from unexposed

mothers, in the presence of toxic cyanobacteria. This effect was mainly elicited in

response to maternal exposure to toxic cyanobacteria early in life and less so during

reproduction. None of these effects were explained by changes in egg size. A meta‐

analysis using our and others’ experiments suggests that the adaptive value of maternal

effects to cyanobacteria exposure is weak at best. We suggest that the beneficial

maternal effect in our study is an example of phenotypic accommodation spanning

generations, rather than a mechanism evolved to transmit information about

cyanobacteria presence between generations.

115

P17

Studying the evolutionary process with genetic algorithms

Edith Invernizzi


Genetic Algorithms (GAs) are a stochastic search algorithm that exploits the

mechanisms of biological evolution -selection, reproduction, mutation and crossover-

to reach optimal solutions in a large, complex fitness landscape where best-performing

solutions cannot be found analytically. Despite having originally been devised as a tool

for the study of the evolutionary process itself, they have had limited applications in

evolutionary biology, mainly in the optimisation of analytical tools (e.g. in

phylogenetics), or in the simulation of complex trait evolution (e.g. cognitive processes)

– to the point that they are mainly used in engineering. Here, I use GAs to study how

differences in the process of selection and reproduction may lead to different end

points in the fitness landscape. I also analyse the dependence of this trajectory on other

elements that are known to have an important role in the biological evolutionary

process: namely, population size and mutation rate. I use the Sir Philip Sidney’s Game,

a game theory problem applied to the study of the evolution of signalling, as a test bed

for our query. I compare five ways of modelling selection and reproduction in the

algorithm and I look at the relative frequencies of Evolutionary Stable Strategies,

Evolutionary stable Sets and other, evolutionary unstable, solutions emerging under

each variant, to analyse the reliance of each method on landscape exploration versus

exploitation of the locally optimal solution. Because differences in the outcome results

from how strictly reproduction depends on quality and on relative reproductive fitness

of the higher to lower-quality solutions, these results have interesting implications for

the effect of different population structures in biological evolution. This study highlights

the usefulness of this simulation tool in improving our understanding of the

evolutionary process itself.

116

P18

The evolutionary origin of our moral beliefs

Man-Him Ip

University of Birmingham, UK

In recent years, philosophers have discussed evolutionary debunking arguments of

morality (EDA) extensively. These arguments basically start with an empirical claim that

evolutionary forces have had a significant influence on what moral beliefs we have.

Evolutionary debunkers then aim to show that the epistemic status of our moral beliefs

is diminished due to that influence.

However, we first need to know at least how evolution has influenced and shaped our

moral beliefs, if it really has done so. I attempt to draw a very rough outline of the

nature of moral beliefs. Basically, there are at least three essential features or key

characteristics of moral beliefs:

(1) There are some core contents of moral beliefs that are commonly shared in most

communities

(2) Moral beliefs are reliably connected to our motivation

(3) Moral beliefs are connected to praise and blame

I argue that a plausible view of the evolutionary origins of our moral beliefs must be

able to explain how our moral beliefs came to have those three essential features. In

this paper, I am going to introduce and evaluate two different theories of how evolution

could have influenced and shaped our moral beliefs. I call these two views the

adaptation account and the exaptation account.

I will explain how these two accounts can provide a plausible explanation of how

evolution could have influenced our moral beliefs. In other words, these accounts can

explain why having moral beliefs with the three essential features is an adaptation and

an exaptation respectively. However, I remain neutral on which of them could be the

best account of the evolutionary influence on our moral beliefs. What I will demonstrate

is that these two accounts have their own plausibility.

117

P19

Genetic assimilation in the fossil record: phenotypic plasticity and accommodation in

Cambrian arthropods

Illiam Jackson, Madeleine Bohlin, Richard Mann, Graham Budd


Genetic assimilation describes a process in which an initially plastic response of an

organism to the environment becomes constitutively expressed through quantitative

genetic change. Here we suggest that the Cambrian Series 3 trilobite-like arthropod

Agnostus pisiformis and subsequent members of its evolutionary lineage allow us to

probe the fossil record for signals of genetic assimilation. The lineage is geographically

broadly represented, as well as stratigraphically well-defined, stretching through a

period of increasing anoxic/euxinic conditions interpreted as stressful to the organisms.

Using elliptical Fourier analysis (EFA), we quantify the morphology of specimens

recovered from geographically and stratigraphically varied assemblages, including

several coeval but geographically disparate localities. For each of these assemblages we

determine relative levels of anoxic/euxinic stress using redox-sensitive metals (Mo, U

and V) as proxies, analysed via mass spectrometry of the sediment.

We uncover an evolutionary pattern in which A. pisiformis and the subsequent member

of the lineage, H. obesus, exhibit directional morphological change as well as increasing

morphological variation in response to increasing anoxic/euxinic stress. The most

recent member of the lineage, T. holmi, responds to the relaxation of this stress by

reducing its variability around a new morphological mean within the range of the

lineage’s earlier expansion. We interpret this pattern as (1) environmental stress

inducing novel phenotypic variation, (2) selection acting on the induced phenotypic

variation followed by (3) canalization of the initially plastic response; in other words,

genetic assimilation.

118

P20

Extended cognitive systems, meaning, and evolutionary transitions

Hilton F Japyassú

Universidade Federal da Bahia, Brazil

Considering that non-biological entities are capable of information processing, and that

cognition is by and large considered as adaptive information processing, the extension

of biological cognition to worldly components is slowly turning into a sound scientific

enquiry. This is relevant for an extended evolutionary synthesis, since worldly

information processing that is tightly coupled to organism functioning can evolve as to

be encapsulated by the organism itself: the organism extends towards specific

components of the world, thus creating larger evolutionary units. Also, since extended

cognition is tightly coupled to developmental learning processes, the new unit is not

random, but rather a fully adjusted new phenotype, thus increasing evolvability.

Extended cognition is thus a promising research program, but one should avoid

important pitfalls that could halt its development. If information is taken as the only

relevant aspect of cognition, with no room for meaning or representations, extended

cognition risks excluding large fields of cognitive enquiry. We build on recent theoretical

developments on biological autonomy to track the emergence of meaning within

dynamical, information processing systems. Meaning requires the generation of

subsystems that are both autonomous and integrative. A cognitive system emerge with

new organisation levels, integrating informationally the performance of previously

disparate subsystems; it requires regular responses to similar external triggers,

contextual adjustment, and internal consistency. We depict successful instances of

extended meaning systems, demonstrating that meaning expansion emerges from self-

organisation within unpredictable environments, with the creation of hybrid brain-

environment loops. It turns out that the extension of cognition to worldly components

works by selecting out the expansion of meaningless information processing, paving the

way to major evolutionary transitions.

119

P21

Mechanisms of hormone mediated parent-offspring conflict

Neeraj Kumar1.3, Martijn van Faassen2, Bonnie de Vries1, Anja Lohrentz3, Ido Kema2,

Manfred Gahr3, Ton GG Groothuis1

1Behavioural Biology, Groningen Institute for Evolutionary Life Sciences, University of

Groningen, The Netherlands; 2Laboratory Medicine, University Medical Center

Groningen, University of Groningen, The Netherlands; 3Behavioural Neurobiology, Max

Planck Institute for Ornithology, Seewiesen, Germany

Vertebrate embryos are exposed to a varying degree of maternal hormones, having

profound consequences for their later phenotype. Bird species have been used

extensively to study these effects, with a strong focus on ecological and evolutionary

factors affecting maternal hormone allocation to the egg, showing systematic variations

among different eggs of the same nest, between-nests, and between-females.

However, the embryo itself is often seen as a passive responder. Its actual role in

adapting to and translating maternal hormonal signals, and its role in the evolution of

parent-offspring conflict, has largely been ignored. By focussing on early hormone

dynamics in the egg and embryo we found that (1) The active maternal hormones in the

egg, such as progesterone and testosterone, are substantially and rapidly metabolized

by the embryo, with a corresponding increase in much less potent metabolites; this

suggests that the embryo itself can time its exposure to the maternal signal by

converting less potent hormones back to potent ones when needed; (2) The dynamics

of androgen metabolism differs systematically between eggs of different laying order;

this opens the possibility that the embryo can deal with the maternal signal depending

on the context it experiences, which may depend on yolk composition and incubation

pattern; (3) The embryo expresses steroid receptors in its extra-embryonic membranes

even before its own hormone production starts, indicating that selection has favoured

the early perception of the maternal signal; and (4) The embryo downregulates these

receptors in response to increased egg steroid levels, indicating that the embryo can

regulate the effectiveness of the maternal signal. Altogether, the results imply that the

embryo is not simply a ‘slave’ to the maternal signals but can potentially play its own

role in parent-offspring conflict, opening new avenues to integrate mechanistic,

developmental and evolutionary approaches to the study of maternal effects.

120

P22

The impact of individual immune experience on niche construction in red flour

beetles, Tribolium castaneum.

Lai Ka Lo, Caroline Müller, Martin Kaltenpoth, Joachim Kurtz

University of Münster, Germany

Adult red flour beetles, Tribolium castaneum produce external secretions containing

high levels of antimicrobial quinones (benzoquinone and hydroquinone), which have

been shown to inhibit the growth of bacteria, yeast and fungi commonly found in flour

and the beetle’s natural environments. As group living animals that share the same

niches, potentially, the secretions from adult beetles could in turn influence the levels

of pathogenic threat that their conspecifics or offspring are exposed to and thus modify

the selective environment of both the niche constructors and recipients. In response to

the sparse experimental tests of the ecology and evolutionary consequences of niche

construction, we here start using the worldwide pest of stored grains, T. castaneum

along with their natural parasite Bacillus thuringiensis tenebrionis (Btt) as a host-

parasite model system for niche construction.

Interestingly, T. castaneum demonstrated immune priming (i.e. increased immune

response against previously encountered microbes) within and across generations. As

the magnitude of animals’ niche construction likely varies with individual experience,

i.e. exposure to environmental stressors, we hypothesise that individual immunological

experiences in T. castaneum might lead to the construction of individualised niches.

In our studies, individual beetles were either left naive, sham-exposed by injection of

PBS or primed via injection of heat-killed vegetative Btt. Using spectrophotometry and

bacterial 16S RT-qPCR, we compare both the quinone levels and microbial loads in the

bodies and flour of adult beetles undergoing different treatments. We will present our

preliminary work on the influence of individual immune experience on niche

construction and our plan to test the evolutionary consequences of individual niche

construction via experimental evolution.

121

P23

Trait-based theory leads to generality across successional systems

Jessica Haghkerdar, Brian McGill, Maria Dornelas


Succession is the turnover in species and/or traits caused by a sudden availability of

unexploited resources. Succession is one of the oldest fields in ecology, but we still do

not have cross-system generality in our models of how succession progresses. If we

want to achieve generality between systems as disparate as gut microbiomes, whale

falls, and tropical forests, we need to be able to compare them empirically and with a

broad taxonomic scope.

Developing a general understanding of succession will entail standardising between

different successional trajectories. The few cross-system succession studies that do

exist are usually in a narrow range of habitat and disturbance types, but taking a trait-

based approach will allow us to compare responses in any variable between

successional systems.

As a starting point for trait-based synthesis, we present a range of hypotheses on how

key traits and life-history attributes will change during succession. These traits span all

taxonomic groups and are relevant to any ecosystem type. While this framework should

apply to any level of organisation, we particularly encourage its use in multitrophic

communities, which have been under-studied, especially in succession.

Traits used in succession studies should reflect reproduction (number and timing of

offspring), performance (how an organism fares in one patch in space and time), and

heterogeneity (how an organism copes with changing conditions). Building on

traditional r-K life history divisions, this framework specifically highlights coping with

changing conditions over time, as in succession. This deliberately cross-system, cross-

taxon, multitrophic framework is suitable for both system-specific studies and

multisystem comparisons. Its future application will help both applied and conceptual

ecologists predict and manage systems undergoing succession and restoration.

122

P24

From anisogamy to intrasexual competition

Ivain Martinossi, Mattias Siljestam


The biological evolution of separate sexes in animals, females that produce eggs and

males that produce sperm, is a fundamental question of evolutionary biology;

furthermore, this initial difference between the sexes is thought to have resulted in a

cascade of evolutionary steps that influenced many aspects of sexual dimorphism,

including the intensity of competition for mating opportunities. Although it has been

advocated that males compete more often and more intensely than females in animals,

and that this pattern finds its roots in the state of anisogamy (gamete size dimorphism),

we suggest that the evolution of anisogamy may not always spur the evolution of male-

bias competition for mating.

Using the method of adaptive dynamics, we develop a model of the coevolution of

gamete size and competition for mating, starting in a population without sexual

dimorphism. We show, in accordance with previous work, that the evolution of

anisogamy is highly sensitive to population density and survival constraints. We also

find that anisogamy is often accompanied by the evolution of a sex-bias in competition

for mating. Finally and most importantly, competition for mating may evolve to be

either female or male-biased, depending on the competition trait considered and which

were the conditions that triggered the evolution of anisogamy in the first place.

123

P25

Diet-based developmental plasticity in a freshwater isopod

Moritz Lürig, Kim Kaltenbach, Blake Matthews

Eawag, Switzerland

Developmental plasticity is common in natural populations and can play an important

role in adaptive evolution. When developmental plasticity is heritable and exhibits

genetic variation in a population, it can evolve when selective environments change.

Evidence for the evolution of plasticity can be obtained by (i) measuring the heritability

of developmentally plastic traits, (ii) identifying the causes of plasticity in natural

environments, and (iii) testing how putative agents of selection affect the distribution

of developmental plasticity over time (within or across generations). The freshwater

isopod, Asellus aquaticus, is a useful animal model to study the evolution of

developmental plasticity in natural populations. Cryptic pigmentation of A. aquaticus is

thought to evolve rapidly in response to predator-mediated selection. However,

pigmentation is a developmentally plastic trait - individuals are born with low

pigmentation, and irreversibly accumulate pigmentation over their lifetime. In a

common garden experiment with 29 families (split-clutch, full-sib design), we show that

the rate of pigmentation accumulation during development varies among families and

increases with a high protein diet. Further, we found that fast growing individuals on

low protein diets have a higher survival probability when they also have higher rates of

pigmentation accumulation. To test whether pigmentation improves crypsis we

performed predation trials with a visually foraging predator (i.e. stickleback fish), in

which we manipulated both the matching background (i.e. dark and light sand) and

frequency of background-matched individuals (i.e. low and high frequency of dark and

light phenotypes). In these experiments, we find no evidence that fish impose either

divergent or frequency-dependent selection on isopod pigmentation. Overall, our

results suggest that natural isopod populations harbour genetic variation in diet-based

developmental plasticity of pigmentation, but the adaptive value of this plasticity is still

unknown.

124

P26

Global human niche construction: an evolutionary trap?

Andra Meneganzin, Francesco Suman, Telmo Pievani

Research Unit in Evolutionary Biology, Department of Biology, University of Padua, Italy

Niche construction theory posits that organisms can act as important agents of

selection, by modifying biotic and abiotic environmental conditions. To test the

presence and the evolutionary impact of niche construction, some criteria have been

identified (Matthews et al 2014): 1- an organism must significantly modify

environmental conditions, 2- organism-mediated environmental modifications must

influence selection pressures on a recipient organism; 3- there must be an evolutionary

response in at least one recipient population caused by the environmental modification.

Homo sapiens has begun, by the late Pleistocene (Boivin et al 2016), in engaging in niche

construction activities that meet those criteria, being however recognizable for some

distinctive features: the rapidity of the processes, their global impact, the strong

ecological inheritance of anthropic environmental modifications. Here we aim in

particular at assessing the evidence for the third criterion, thus providing arguments for

alterations in evolutionary trajectories due to anthropic activities. Moreover, human

modifications have often brought to significant deleterious environmental

consequences: it has been established that we’re losing ~11,000 to 58,000 species

annually, changing ecosystem functions and services in a dramatic way (Dirzo et al

2014); by altering the climate, we have altered chemical composition of water, species

distribution and their habitat. We discuss the extent to which global warming and

biodiversity loss fall within the definition of niche construction activities. In fact, niche

construction is often referred to as an adaptive process and global warming and

biodiversity loss can indeed be intended as a by-product of adaptive human activity in

the short term. However, this same activity might not result an adaptive one in the long

term, as impoverished ecosystems might feedback negatively on human prosperity.

Therefore, human niche construction so described might take the form of an

evolutionary trap for Homo sapiens itself, resulting in a no longer adaptive process.

125

P27

A weapons-testes tradeoff in males extends to female traits

Christine W Miller, Paul N Joseph, Rebecca M Kilner, Zachary Emberts

University of Florida, USA

Trade‐offs among costly fitness traits are a central tenet of evolutionary theory. An

organism cannot optimally invest in all traits associated with reproduction,

development and survival, but must balance competing functions. Despite decades of

research on such life history tradeoffs, we still know remarkably little about what

governs resource allocation decisions. In this experimental study, we compared trait

size in males and females and used the phenomenon of autotomy (dropping of limbs)

as a form of phenotypic engineering to reveal allocation patterns. In Narnia femorata

(Hemiptera: Coreidae) males use their hind limbs as weapons in male-male contests

over access to females, while females use their hind limbs only for locomotion. We

induced hind-limb autotomy in a sample of both males and females and measured

resulting gonadal size to test the extent to which tradeoffs between sexually-selected

weapons and testes in males are found in the homologous traits in females. As

expected, insects that lost a hind limb during development grew larger gonads. Further,

when we adjusted for physical differences in size, female ovaries increased more than

male testes following autotomy. Thus, a documented weapon-testes tradeoff in males

is also a limb-ovary tradeoff in females, a similarity that was not expected. These results

suggest current theory on resource allocation may place an overemphasis on trait roles,

while resource allocation patterns may more closely follow tissue type and expense of

traits.

126

P28

Assessing adaptive and non-adaptive evolution of the glycolytic enzyme

triosephosphate isomerase

Ricardo Muñiz Trejo, Sergio Romero Romero, Daniel Alejando Fernández Velasco

Universidad Nacional Autónoma de México, Mexico

Protein evolution studies have been dominated by accounts that explain

physicochemical phenomena such as reversibility, folding, stability, oligomeric state,

etc., as adaptive features, aligning with the tenets of Standard Evolutionary Theory.

However, very few of them have inquired into the validity of these hypotheses from a

broader evolutionary perspective. Understanding the underlying evolutionary

processes in biochemistry might provide us with a better way to explain how evolution

shapes the properties of macromolecules and how these properties set evolutionary

pathways. In this work, we are interested in exploring which evolutionary processes,

including those highlighted by the Extended Evolutionary Synthesis, are changing (or

not) the thermostability of the glycolytic enzyme triosephosphate isomerase (TIM) in

thermophilic archaea. For this enzyme, a change in the oligomeric state, from a dimer

to a tetramer, is assumed to be the adaptation to life at high temperatures in Archaea.

We are characterizing recombinant TIMs from archaea of the order Methanococcales

with different growth temperatures (38 - 85 °C). Our experiments show that both

tetrameric state and high thermostability (>90 °C) are preserved in TIMs from

mesophilic representatives. Furthermore, hyperthermophilic representatives retain

high catalytic activity even at 25 °C. These results suggest that there are some

phylogenetic constraints involved in the evolution of TIMs from this order that are

keeping these enzymes with high thermostability. Further studies regarding the

thermostability of other proteins from these organisms need to be performed, but our

results are coherent with other works that propose a readaptation of thermophilic

Methanococcales, since thermostable proteins may contribute to the evolvability of this

clade towards life at high temperatures.

127

P29

The evolution of hierarchical structure: separating the causes and consequences of

evolvability

Frederick J Nash, Loizos Kounios, Kostas Kouvaris, Richard Watson, Danesh Tarapore


It has previously been suggested that the evolutionary causes of hierarchical

organisation in biological systems are different from the evolutionary consequences of

that hierarchical organisation. Specifically, it has been shown that evolved hierarchy can

have the long-term consequence of facilitating evolvability, but that the short-term

selective pressures that cause hierarchy to evolve can be a side effect of selection in

systems where regulatory connections are costly. Here we investigate the conditions

for the evolution of hierarchy and its relationship to evolvability in the context of

development controlled with a gene regulation network.

Such networks can exhibit non-modular structures, modular but not hierarchical

structures, and modular structures with internal hierarchy. Using this model, we

identify conditions under which hierarchical structures are favoured by short-term

selection before any impact on evolvability may be observed. We rule-out the long-term

advantages of evolvability as a necessary cause, and present cases where hierarchy

evolves in static selective environments where no evolvability benefit can be observed.

These results show that the distinction between the short-term selective causes of

hierarchy and the long-term consequences of hierarchy can be much more distinct than

previously documented, and can even be separated completely. We describe

experimental modifications where hierarchy facilitates evolvability but does not evolve

and cases where hierarchy evolves without facilitating evolvability.

We discuss what would be required to determine whether the occurrence of hierarchy

in nature evolved for evolvability or for other reasons, and whether there might be a

deeper reason why short-term and long-term benefits might be coincident in natural

systems.

128

P30

Spandrels and trait delimitation: no such thing as “architectural constraint”

Mark E Olson

Instituto de Biología, Universidad Nacional Autónoma de México, Mexico

40 years ago, Gould and Lewontin used the metaphor of a building’s “spandrels” to

highlight that organismal traits could be the inevitable consequence of organismal

construction, with no alternative configurations possible. Because adaptation by

natural selection requires variation, regarding a trait incapable of variation as an

adaptation could be a serious error. Gould and Lewontin’s exhortation spurred

biologists’ efforts to investigate biases and limiations in development in their studies of

adaptation, a major methodological advance. But in terms of the metaphor itself, over

the past 40 years there are virtually no examples of “spandrels” in the primary

literature. Moreover, multiple serious confusions in the metaphor have been identified

and clarified, for example that the “spandrels” of San Marco are pendentives, and

pendentives are perfect examples of adaptation. I look back over the sparse empirical

fruits of the “spandrels” metaphor, and ask what the clarifications of the past 40 years

mean for biological theory and practice. I conclude that if there is anything to be

rescued from the clarified spandrels metaphor, it not “constraint” at all. Instead, it is

the still-unresolved issue of trait delimitation, which is how to parse organisms into

subsets that are tractable and biologically appropriate for study.

129

P31

With a little help from my friends: the role of the microbiota in dung beetle

diversification

Erik S Parker, Daniel B Schwab, Armin P Moczek

Indiana University - Bloomington, USA

Understanding the processes that enable and shape the generation of biodiversity is a

major objective of evolutionary biology. Traditionally, explanations have focused on the

role of divergent selective pressures acting on generations of individuals. However,

increasing appreciation for the vital role of host-associated microbial symbionts has

added complexity to this notion of individuality, and raised the possibility that selection

may occasionally, or perhaps frequently, act on teams of cooperating taxa rather than

individuals. Yet few study systems exist where such perspectives can be assessed

experimentally. My research aims to address this shortcoming by exploring the role gut

symbionts play in the diversification of their dung beetle hosts. Despite feeding on

nutritionally limited mammal dung throughout all stages of their life, Onthophagus

dung beetles have been extraordinarily successful both ecologically and evolutionarily

as measured by their expansion onto every continent except Antarctica, specialization

onto an enormous array of dung types, and speciation into well over 2000 extant

species. Using a combination of microbiota removal and transplant experiments we

have shown that Onthophagus larvae inherit their gut microbiota vertically through a

maternal fecal deposit known as a pedestal, that pedestal-derived microbiota enhance

larval growth and survival especially under stress as well as protect against pathogens,

and that different beetle host species have specialized onto different and functionally

non-equivalent microbiota. Current efforts utilize recently established exotic dung

beetle populations to assess host microbiota fidelity during the colonization of novel

habitats as well as the role of microbiota in limiting, enabling, or biasing host range

expansions. My talk will explore the most recent findings of this research area and their

larger implications for understanding the role of host-microbiota interactions in the

generation of biological diversity.

130

P32

Transposable elements: the unexpected junk that shaped genomes and challenged

evolutionary biology

Valentina Peona


Barbara McClintock first described transposable elements in 1951 but her discovery

remained ignored for decades, probably because her system of ""controlling

elements"" did not fit any known genetic model although they shared some conceptual

similarities with the (already at the time well-known) operons.

In addition to be ignored, in the 1980 two important reviews published in the same

Nature issue labelled and stigmatized repetitive DNA as junk DNA with no phenotypical

relevance even though transposable elements themselves have been discovered

because of their phenotypical impact. This label is still in use but thanks to scientific and

technological advancement of the last decades, we have been able to appreciate them

as a great substrate for evolution by molecular tinkering.

Transposable elements changed our view of genes and genomes, from static entities to

plastic and fluid. In fact, some transposable elements have been co-opted as

telomerases, for the acquired immune system and placental evolution/development.

They are inevitable genomic parasites from which genomes must be protected but also,

in time, became essential components of eukaryotic genomes.

Nowadays, we know that transposable elements regulate gene expression, play an

active role during embryogenesis, provide neuronal plasticity and they also found use

in the biotechnology industry as gene editing tools and certainly more roles and

functionalities are waiting to be discovered.

In this study I investigate the historic context in which Barbara McClintock discovery

occurred, how the Modern Synthesis framework at that time influenced the reception

and interpretation of her work, how transposable elements still challenge our

comprehension of genome evolution (e.g. casting questions about what is actually

neutral or under selection).

131

P33

Weismann, Crick, and others: flows of information in evolution

Arnaud Pocheville

CNRS, and Université Paul Sabatier, France

In previous work I have argued that one classical principled reason to separate

development and evolution is that they are supposed to concern different levels and

time-scales. I have also pointed to new theoretical approaches that aim at questioning

this separation.

In this talk I will discuss another kind of principled reason for this separation, one

pertaining to biological information. Classically, development has been supposed to

deal with the expression of existing information (mostly genetic information). This

simplifies development to a one-way flow of information (from genes to the

environment) at the individual level. Evolution, on the other hand, especially by natural

selection, is supposed to deal with the inscription of new information. This simplifies

evolution to another one-way flow of information (from the environment to the genes),

but now at the population level.

I will mention why the separation of flows of information can be attractive and briefly

connect it to the question of the separation of time-scales.

132

P34

Delineating the boundaries of adaptation by tracing histories of environmental

induction

Ahva L Potticary, Erin S Morrison, Alexander V Badyaev


Adaptations change by reconfiguration of their components or by modification of

homeostatic systems that maintained stability of preceding states. However,

comparison of extant adaptations offers only indirect insight into the relative

importance of these processes in evolution. Here, we overcome this limitation by

directly tracing coevolution of adaptation components (diverse carotenoids taken up by

growing ornamental feathers), with adaptation boundaries (sensitivity of

developmental decision to forgo the feathers’ structural differentiation that limits

accumulation of carotenoids). We found that along known historical trajectories linking

native and 48 recently established populations, evolutionary changes in developmental

boundaries of adaptation were not specific to individual carotenoids, but instead

reflected the extent to which carotenoids were integrated into organismal processes

and were environmentally predictable. Across all historical sequences, uptake of

internalized carotenoids did not change the boundaries of feather developmental

response, instead modifying the rate and precision of the response. In contrast,

accommodation of uncommon dietary carotenoids strongly modified both the rate and

the pattern of developmental response, especially in new populations. We found that

largely inductive effects of carotenoid uptake on feather differentiation in younger

populations were progressively converted into anticipatory effects as populations

persisted in local environments, and show that this was due to increasing organismal

coordination of feather differentiation decisions. These findings strongly implicate

cooption and modification of generalized stress buffering mechanisms that shield

feather growth and differentiation (i.e., adaptation boundary) as a starting point for the

evolution of greater expression of external pigments within intricately structured

feathers.

133

P35

The interaction of learning and evolution can enable adaptive behaviours that are not

presupposed in the learning ability

David Prosser, Alfredo Rago, Markus Brede, Richard A Watson


Learning is a highly adaptive form of phenotypic plasticity where a behaviour can

change over an individual’s lifetime as a result of experience. Innate behaviours can also

be adaptive but do not change over an individual’s lifetime. In the absence of a

mechanism for the inheritance of acquired characteristics, learned behaviours cannot

influence innate behaviours directly. Nonetheless, the genetic assimilation of learned

behaviours, often described as the Baldwin Effect, enables learned behaviours to guide

the genetic evolution of innate behaviours indirectly but in a systematic manner.

Previous models of this effect have, however, presupposed the ability of lifetime

learning to produce appropriate adaptive behaviours. This makes a ‘plasticity-first’

concept of evolution based on such models open to the criticism that, although genetic

evolution may follow behind a high-fitness acquired behaviour, the ability to produce

the high-fitness behaviour in the first place was presumably the result of prior genetic

evolution. Here we use a computational model to study the interaction of learning and

genetic evolution in a multi-peaked fitness landscape where high-fitness behaviours

cannot be quickly and reliably discovered by the learning process alone (nor by genetic

evolution alone). We find conditions where the effect of genetic evolution is to

incrementally reduce the dimensionality of the behavioural space that learning

explores, not simply to assimilate the solutions that learning has already found, but in

a way that causes the learning process to find higher fitness behaviours more reliably

than was previously possible. This bidirectional interaction between evolution and

learning is a form of plasticity first evolution capable of producing adaptations that are

neither presupposed in the learning ability nor provided by prior genetic evolution.

134

P36

Naturalising the language faculty: the role of structures and developmental processes

Giuliana Pulvirenti, Salvatore Ivan Amato, Alessandra Falzone

University of Messina, Italy

The form/function dichotomy is one of the oldest and conceptually most pregnant

distinction made in order to investigate the nature of organismal traits. Although the

use of these two fundamental concepts is widespread in the field of contemporary

evolutionary biology, their application is not so straightforward and can be theoretically

problematic depending on the level of analysis considered (Love 2007). Moreover, in

recent years this topic has gained a larger amount of attention due to insights obtained

from Evo/Devo approaches (Minelli 2003; West-Eberhard 2003). The overmentioned

distinction is relevant also among cognitive strains of research that ought to describe

and explain in evolutionary terms complex and species-specific behaviours adopting the

comparative method. The case of language faculty is, in this respect, paradigmatic. The

present work aims at understanding language as a complex biological function while

defending the necessity to pay attention to structural elements and ontogenetic

processes to properly characterize it and analyse it from an evolutionary point of view.

Most of the hypothesis formulated in this area suffer of a functionalist bias, often

neglecting the role that morphological structures as well as ontogenetic processes exert

on the emergence, development and evolution of animal behaviors, thus offering at

best only a partial view. The interplay of these structural elements is modulated in and

by the socio-ecological context in which the organism is to live in, constraining, while at

the same time providing, a range of functional possibilities. That is to say functions are

both epistemologically and chronologically dependent on forms and structures. This

aspect has been heavily recognised among new embodied approaches to cognition

(Shapiro 2004), in contrast to genetic and cerebrocentric/computational ways of

thinking about human cognitive functions (Berwick & Chomsky 2016).

135

P37

Teleology and evolution

Rasmus Sandnes Haukedal

Durham University, UK

In Organisms, Agency, and Evolution (2015), D. M. Walsh argues that a new synthesis –

which goes beyond the Modern Synthesis (MS) – cannot merely be about extending the

scope of what evolution is about but must thoroughly reformulate its main tenets. It is

thus wrong to assume that the basic framework of the MS could be left intact by a new

model. We must formulate a comprehensive new theory, in which history is

reintroduced into evolutionary thinking – not merely add factors with evolutionary

significance to a pre-existing framework.

In Walsh’s view, what is at stake in this reorientation is defining what counts as

‘evolutionary’ in the first place. He proposes an ‘a posteriori’ approach, in which

evolutionary significant factors are not given but established retroactively. This means

that such factors are transhistorical: while occurring at a specific historical point, they

change the trajectory and dynamics of evolution. Furthermore, Walsh states that a new

evolutionary synthesis demands the restoration of some sort of teleology – which he

associates with phenotypic plasticity. We find the same idea in Scott Turner’s Purpose

& Desire (2017), where it is also linked up with homeostasis and self-organization. They

both argue that teleological explanations are indispensable to biology. I will examine

what this move entails and apply it as an analytic tool to judge different attempts at

transgressing the MS framework. Evidently, rethinking individuality is required, as is the

inclusion of more factors than genetics. It is, however, not clear how Walsh’s ‘situated

Darwinism’ differs from other attempts at going beyond MS, and whether these

proposals are complementary or not. They all share an emphasis on the developmental

effects on evolution, linked up with the question of how variation is produced. It seems

the question of teleology might ultimately be what separates them.

136

P38

The evolution of adaptive epigenetic inheritance

Zuzana Sekajová, Elena Rosa, Irja Ida Ratikainen, Martin Lind


Environmental heterogeneity can influence the evolution of inheritance systems. If the

environment is stable, genetic specialization should be favored by selection. If the

environment is slowly fluctuating (cycle length spanning several generations), then the

parental environment represents a reliable cue to the offspring and epigenetic

inheritance is predicted to evolve. In the contrast, if the environment is fluctuating fast

and unpredictably, there is no correlation between parental and offspring environment

and phenotypic plasticity, bet hedging and/or generalism are predicted to be adaptive.

My overall aim is to investigate the role of environmental heterogeneity for the

evolution of inheritance systems with special focus on epigenetic inheritance. I will test

this using experimental evolution in nematode worm Caenorhabditis remanei by

manipulating temperature environment. Worms adapted to 20°C were selected in four

regimes differing in cycle length for 113 non-overlapping generations: Warm with

constant 25°C, where 25°C is a novel and mildly stressful environment, Cold with

constant 20°C, Cyclic where temperature cycles between 20°C and 25°C every 8

generations and Random where temperature fluctuates randomly between 20°C and

25°C.

I will conduct trans-generational phenotypic assays in a fully factorial design (parental

temperature x offspring temperature) in which I will test worms for the key life history

traits as reproduction and growth. I predict the evolution of epigenetic inheritance in

worms from cyclic environments, specialization in worms from constant environments

and the evolution of plasticity/bet hedging in worms evolving in random environment.

In addition, I will investigate the molecular basis and regulation of plastic, epigenetic

and genetic trait induction by small RNA sequencing which is the main epigenetic

mechanism in the nematode worms.

137

P39

Unique evolutionary properties of plastic traits: an empirical study of erosion of

phenotypic plasticity under relaxed selection

Helen Spence-Jones, Michael M Webster, Kevin N Laland


While there are many hypotheses concerning the dynamics of plasticity which have

proof-of-concept (Baldwin effect, plasticity-first, plasticity-mediated persistence etc),

few of them have been found to be universal rules. I suggest that this is because

plasticity is such a broad topic; dynamics are specific to subtypes of plasticity. Predicting

the evolutionary dynamics of a plastic trait, and the applicability of hypotheses

concerning this, requires understanding of when and how the underlying mechanisms

of plasticity affect trait evolution. While many aspects of plasticity are only quantitively

different from non-plastic traits, there are some unique properties - generally to do with

decoupling of genotypic & phenotypic variation and the development of novel traits.

These unique properties of phenotypic plasticity come together in the response of a

population to a novel environment (one which has been under relaxed selection): here

there is the potential for extrapolation of reaction norms to generate novel traits,

and/or release of previously-selection-shadowed cryptic genetic variation. Here I

present the results from my research into erosion of salinity-based plasticity under

relaxed selection in threespine stickleback populations, in an attempt to investigate the

dynamics of plastic traits under relaxed selection and potentially link this to the

mechanism of plasticity of the traits concerned. I show that trait plasticity does appear

to erode under relaxed selection, but that patterns of erosion are not consistent

between populations.

138

P40

A culturally driven plasticity hypothesis for the evolution of human life course

Francesco Suman

Research Unit in Evolutionary Biology, University of Padua, Italy

Homo sapiens' life history pattern possesses both fast and slow components, in a

combination that is unique among the extant great apes. The role played by phenotypic

plasticity as a non-genetic means of adaptation to evolutionary challenges is still

debated today. While life history parameters are shaped at a species level by genetic

adaptations via natural selection, they remain very sensitive to changes in the

environment. Relying on evidence from primates, extinct hominins and extant humans

and on key explanatory tools of the Extended Evolutionary Synthesis (phenotypic

plasticity, niche construction and inclusive inheritance), it is likely that environmentally

induced plasticity led the way in human life history evolution, promoting subsequent

genetic accommodation (Kuzawa and Bragg, 2012). It will be shown that the increase in

brain size (modified brain growth and developmental rates) in hominin evolution was

dependent on the appearance of certain life-history traits, which are in turn dependent

on modified extrinsic mortality rates (Robson and Wood, 2008; Hublin et al., 2015). To

the extent that culturally transmitted behaviors altered the selective pressures that

acted on life history traits across generations during human evolution, a culturally

driven plasticity dynamic shaping human life history traits can be identified. Two case

studies in particular will be discussed: the human adaptations to the domestication of

fire and the self-domestication hypothesis. The relationship between a relaxation of the

selective regime and the emergence of plastic responses to changing environment will

be also discussed. Finally, given this activity of cultural mitigation of selection altering

environmental mortality rates is powered by an enlarged and reorganized brain, it is

argued that the causal relationships in play in this evolutionary dynamic is to be

conceived as reciprocal rather than linear (Laland et al., 2015).

139

P41

Unique, essential, and unknown: the enigma of taxonomically restricted, essential

genes of unknown function

Change Tan, Andrew Jones, PA Nelson

Department of Biology, University of Missouri, USA

The advent of automated DNA sequencing in the mid-1990s enabled the development

shortly thereafter of transposon mutagenesis screens of entire genomes, to determine

what genes (and their protein or functional RNA products) were essential for cell

viability under laboratory conditions. Two decades later, these experiments, as well as

systematic or targeted gene deletion experiments, have consistently returned an

unexpected result: many essential genes are (1) taxonomically restricted in their

distribution, not universally, or even widely, shared, but also (2) when annotated, these

genes are most often classified as "unknown function." Evident already in the initial

transposon mutagenesis screens of Mycoplasma genitalium (Hutchison et al. 1999),

where approximately one third of the essential genes were listed as "unknown

function," the same "unique, essential, and unknown function" signal has been found

in all three domains of life, e.g., in the proteobacterium Caulobacter crescentus

(Christen et al. 2011), the archaean Sulfolobus islandicus (Zhang et al. 2018), and the

eukaryote Drosophila melanogaster (Chen et al. 2010). We synthesize the findings of

all whole genome transposon mutagenesis and targeted deletion experiments to date,

with respect to their fraction of "unique, essential, and unknown" sequences, and

consider the theoretical consequences of this pattern for theories of cell function and

evolution.

140

P42

The evolution of relatedness and evolutionary transitions in individuality

Christoph Thies, Markus Brede, Frederick J Nash, Richard A Watson


Fraternal transitions in individuality, such as the transition from unicellular to

multicellular organisms, are central to the evolution of complex life. Such a transition

involves and depends on an increase in the genetic relatedness of the particles within

collectives, such as cells within organisms. Most models of social evolution take

relatedness to be a parameter defined by exogenous factors. However, this assumption

precludes analysis of how this crucial population property arises as a product of the

evolutionary process on the particle level. For such an analysis, relatedness must be

endogenised. Although some previous models have studied mechanisms, such as the

evolution of group size or inbreeding, where relatedness is modified by individual traits

under selection, general conditions for the evolution of relatedness have not been

identified. Here we show that no net change in relatedness occurs when the interaction

between particles is described as additive pairwise game with two strategies, i.e. a

game without synergistic effects such as a standard Prisoner’s Dilemma. In additive

games, selection for beneficial organisation has opposite effects for each of the two

strategies. While this result requires some assumptions, it illustrates the intuitive idea

that a stable organisation of particles in collectives requires the alignment of benefits

from that organisation. We show that the synergistic effects between particle types in

non-additive games, such as coordination/anti-coordination games, cause such an

alignment and facilitate the evolution of relatedness in a population. Focusing on anti-

coordination games as models of functional complementarity between particle traits,

we study how plastic trait expression evolves in concert with relatedness. This allows

us to identify conditions under which particles organise into the genetically

homogeneous and phenotypically heterogeneous collectives that are the characteristic

consequence of a fraternal transition in individuality.

141

P43

Deconstructing reef complexity: relating coral diversity, rugosity and fractal

dimension

Damaris Torres-Pulliza, Maria Dornelas, Joshua S Madin

Macquarie University, Australia

The relationship between habitat structural complexity and biodiversity is foundational

to understanding community ecology and informing conservation management

strategies. For habitat engineering organisms, like the reef-building corals, the direction

of the complexity-biodiversity relationship is not clear, because the organisms that

create structural complexity often require complexity to settle as larvae. Meanwhile,

studies looking at complexity-biodiversity relationships typically revolve around one

metric, rugosity, and the assumption is that higher levels of rugosity lead to higher

abundances and richness of reef organisms. Here, we test the strength of complexity-

biodiversity relationships using two metrics that capture different aspects of 3D

complexity (rugosity and fractal dimension) as well as explore reciprocal causation of

the relationships. Metrics of structural complexity were quantified from

photogrammetric surveys at 21 reef sites encircling Lizard Island in the Great Barrier

Reef. We assessed the strengths of relationships between these two metrics in 336

two-by-two metre reef plots and the abundance and richness of coral species in the

same plots. The results are nuanced: structural complexity indeed constructs niches for

coral species at the same time as specific species growth forms drive habitat structural

complexity. Rugosity and fractal dimension captured different elements of the reef

complexity and produced contrasting associations with coral species richness and

abundance. We discuss the implications for niche diversity and the expected size

distributions of reef-associated organisms, like the reef fishes. A better understanding

of the role of reef structural complexity on reef biodiversity is important to guide

management efforts as reefs continue to rapidly change.

142

P44

Cultural evolution and neuroscience: a pathway to integration

Ryutaro Uchiyama, Michael Muthukrishna

London School of Economics and Political Science, UK

Humans have a remarkable capacity to accumulate adaptive behaviours in the form of

cumulative cultural knowledge, and to transmit these traits across generations in

parallel with genetic transmission. This notion of culture as a second system of

inheritance has become increasingly influential in recent years, and is frequently

proposed as an explanation of our ecological success. The usual subtext is that whereas

genetic information is stored in DNA, cultural information is stored within brains and

the machinery of high-fidelity social learning enables this information to be replicated

across brains. While this picture is not incorrect, it fails to capture much of what makes

the human brain–culture interface so effective as an adaptive mechanism. In order to

rectify this, we integrate evidence from neuroscience and cultural evolution, and lay

out the following series of arguments:

(1) Although neural plasticity is usually discussed in the context of how the nervous

system responds to deviations from normative input, we should instead view it in terms

of the space of neurophenotypic variation that it enables. (2) Culture exploits neural

plasticity to explore this neurophenotypic space. (3) In most species, neurophenotypic

variation is driven predominantly by environmental variation (via learning), but the

intrinsic evolutionary dynamics of cumulative culture allow it to explore

neurophenotypes not only in the absence of environmental change, but also beyond

the range of variation that could plausibly be achieved by environmental input. Culture

thus realizes functional fit between brains and environments in an indirect manner, by

serving as a rapidly evolvable surrogate of environmental input. (4) In order to

understand human brain function, we need to look at extant cultural variation as well

as cultural evolutionary dynamics.

143

P45

Lost (and found) in translation: circulation of EES-related concepts in two scientific

communities in Mexico

Francisco Vergara-Silva, Alejandro Fábregas-Tejeda

Instituto de Biología, Universidad Nacional Autónoma de México, Mexico

Academic institutions in the United States of America (USA) concentrate most of the

criticism to the Extended Evolutionary Synthesis (EES) from the Standard Evolutionary

Theory (SET). Given that the hegemonic version of biological evolutionism practiced and

taught in the rest of the American continent –including Canada and the assemblage of

(mostly) Spanish- and Portuguese-speaking countries usually identified as ‘Latin

America’– is deeply influenced by SET-aligned research from the USA, inquiry into the

reception of the core proposals of the EES outside of that country is of potential interest

to evaluate the standing of alternative evolutionary perspectives at local/regional and

international scales. Informed by debates on the ‘circulation of scientific knowledge’

held mostly in history of science circles, in this work we present a comparison of

responses from two research/teaching communities in Mexico to the central tenets of

the EES. The two communities addressed here correspond to separate academic

subdisciplines with legitimate links to evolutionary theory –namely, biological

anthropology and (as expected) evolutionary biology. Although in both cases we

identify a polarization of positions –i.e., researchers place themselves in favor or against

the EES in both Mexican communities– we identify that EES-related ideas have

circulated with less resistance and/or have been received more sympathetically by

biological anthropologists than by evolutionary biologists. We discuss epistemic and

sociological explanations for these results, ranging from the entrenchment of

adaptationist/selectionist thinking and the hardening of population genomics (both in

‘acclimatized’ versions) to the power structure in institutionalized life sciences in

Mexico, where prominent SET-sympathetic evolutionary biologists have held high-level

positions for decades. Finally, we argue that ethnography-laden, locally focused

historiography of science studies could foster collaborations with EES-oriented scholars

–including practicing evolutionists, as well as historians, philosophers and sociologists

of science– currently working in traditional ‘metropolitan’ centers of knowledge

production relevant to contemporary evolutionism.

144

P46

Testing the flexible stem hypothesis in stickleback fish: is ancestral morphological

plasticity adaptive?

Matthew A Wund, Nirmiti Borkhetaria, Rajath Kenath

Department of Biology, The College of New Jersey, USA

One way in which phenotypic plasticity might play a constructive role in evolutionary

processes is by influencing the pattern of phenotypic diversification during adaptive

radiation (the “flexible stem hypothesis,” sensu West-Eberhard). If an ancestral stem

group repeatedly colonizes alternative environments, then phenotypic plasticity

repeatedly produces alternative phenotypes, which can then be refined by natural

selection. This scenario would contribute to the evolution of parallel ecotypic variation

within an adaptive radiation. The threespine stickleback radiation is ideally suited for

testing the flexible stem hypothesis because we can compare patterns of phenotypic

plasticity in the extant, oceanic ancestor to the phenotypes of derived, freshwater

ecotypes. We have previously documented patterns of ancestral morphological

plasticity consistent with the flexible stem hypothesis. Our goal in this experiment is to

determine whether that plasticity confers an advantage in simulated open and shallow

water environments, which would further support the hypothesis that ancestral

patterns of plasticity biased the repeated evolution of limnetic and benthic ecotypes.

We reared oceanic stickleback in simulated open and shallow water environments,

respectively. After five months, we tagged and weighed all fish, switched half of the

individuals into the alternate treatment, and reduced rations to promote competition.

After seven weeks under reduced rations, we again weighed all individuals, predicting

that fish who remained in their original treatment would outcompete those with a

mismatched morphology. We found general support for our hypothesis: fish who

remained in their original developmental environment grew more than those who were

switched to the alternative environment, and we could identify specific differences in

body shape that were most successful under each condition. Finally, fish that moved

to the alternative environment continued to exhibit plasticity, such that after five

months, they closely resembled fish that had developed exclusively in those

environments.

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Quantifying niche constructing traits in reef corals

Kyle JA Zawada, Joshua S Madin, Maria Dornelas


Morphology constrains how all organisms interact with their environment and other

organisms, conditioning the environmental and biological contexts in which they are

successful. For autogenic ecosystem engineers such as reef corals, morphological traits

determine not only their fitness, but also the ways in which they modify the

environment. Hence, morphology determines niche construction processes. Here, we

identify quantitative traits that capture the three main axes of variation in morphology

of reef building corals: colony volume compactness, surface area complexity and top-

heaviness. We posit how variation in coral colony morphology translates to variation in

ecosystem function and niche construction processes important for both corals and

reef-associated taxa. These morphological traits can be used to test hypotheses about

the ecological and evolutionary effects of coral niche construction.

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Two senses of biological possibility

Jason Zinser

University of Wisconsin - Stevens Point, USA

Modal concepts are often employed in the evolutionary biological literature despite the

lack of analysis of how these concepts function in the biological domain. What does it

mean to say, for example, that something is biologically possible or impossible? Is there

a sense of modality that is unique to biology? I argue that there are at least two senses

of possibility in biology: a broad sense and a restricted sense. Dennett (1995) described

a broad sense biological possibility where to be possible is to be genetically accessible:

if X is possible, then there is a genotype that codes for X. Among other things, this sense

is not epistemically fruitful (and Dennett, it seems, ultimately agrees). Fortunately, I

argue that there is a second, restricted sense of biological possibility that is both

metaphysically grounded and interesting. This restricted sense ties possibility to the

variation potential of actual or historical lineages. I will rely on the homeostatic

property cluster kind conception of essentialism in order to ground lineages to a set of

essential property clusters. Once lineages have an essence, then possibility can be

restricted to what variation is possible in that lineage. In other words, it becomes

empirically interesting what variations are restricted to a particular lineage (e.g., how

developmental constraints restrict variation). I will defend this view against challenges,

for example Forber (2009), that argue that biological possibility is circumscribed by

evolutionary forces, such as natural selection and drift. Finally, I will gesture toward the

implications of this restricted sense of possibility for other philosophical disputes, such

as biological explanations, evolvability, and novelty.

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Plasticity in mouthpart length in response to developmental diet in leaf-footed bugs

Sara Zlotnik, Pablo E Allen, Christine W Miller

University of Florida, USA

Invasive species have become increasingly common worldwide. One reason why some

species become invasive while others cannot may be their capacity for developmental

plasticity. Species with high plasticity may be able to tolerate a range of novel

environmental conditions, including new dietary challenges. We are beginning a series

of projects to examine the role of developmental plasticity in the invasion process.

Here, we report our findings on the first project, where we examine developmental

plasticity in mouthpart length in the invasive leaf-footed bug, Leptoglossus zonatus

(Hemiptera: Coreidae). This insect feeds on an astounding array of plants from multiple

families and may grow longer or shorter mouthparts during development to make the

most of the available food type. Such a capacity may contribute to its success as an

invader. To investigate this diet-induced plasticity, we raised insects on multiple food

types and measured their resulting mouthpart lengths. Our study provides insight into

how invasive animals cope with dynamic environmental conditions, such as changing

food availability, while colonizing and spreading through their nonnative range.

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Evolution and development of polyphenisms: a mechanistic model based on bistable

switches

Roman Zug, Tobias Uller


Polyphenisms represent a form of adaptive phenotypic plasticity that gives rise to

discrete alternative phenotypes, based on a developmental switch. However, existing

models for the evolution of polyphenisms do not consider the role of development and

therefore cannot explain how the switching behavior emerges and evolves. Our aim

here is to reveal how polyphenisms evolve by modeling the evolution of their underlying

developmental regulatory architecture. In particular, we hypothesize that

polyphenisms are based on bistable switches in the underlying gene regulatory network

(GRN), which usually requires positive feedback and ultrasensitivity. To test this

hypothesis, we develop a theoretical mechanistic model that focuses on the evolution

of the GRN responsible for the polyphenism switch. Using individual-based simulations,

we explore the conditions (in terms of timescale and predictability of environmental

variation) under which both positive feedback and ultrasensitivity emerge in the GRN,

thus giving rise to a polyphenism switch. We will also present preliminary findings on

the implications of the regulatory architecture for the evolution of polyphenisms. Our

model can be applied to a wide range of polyphenisms and will eventually allow us to

address critical yet unexplored questions, and make testable predictions, about the

evolution of polyphenisms and how it is contingent upon underlying regulatory

conditions.

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