intrinsic inter and intraspecific competition of parasitoids-ssnaik tnau
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CENTRE FOR PLANT PROTECTION STUDIESDEPARTMENT OF AGRICULTURAL ENTOMOLOGY
TAMIL NADU AGRICULTURAL UNIVERSITY COIMBATORE, TAMIL NADU-641 003
SSNAIK, TNAU
Chairman Dr. S. Sridharan, Ph. D., Professor of Agrl. Entomology
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INTRINSIC-INTER AND INTRASPECIFIC COMPETITION IN HOST – WASP PARASITOID RELATIONSHIP
DOCTORAL CREDIT SEMINAR – I (0+1)
Chairperson: Dr. S. Sridharan, Ph.D., Professor, Dept. of Agrl. Entomology, CPPS, TNAU, CBE-3.
ADVISORY COMMITTEE
Members:
ScholarS. Srinivasnaik, II Ph.DID.No: 2015800506
Dr.K.Bhuvaneswari, Ph.D.,Professor,Dept. of Agrl. Entomology,CPPS, TNAU, CBE-3.Dr.S.Mohan Kumar, Ph.D.,Professor,CPMB&B,TNAU, CBE-3.Dr.S.Nakkeeran, Ph.D.,Professor, Dept. of Plant Pathology,TNAU, CBE-3.Dr. S.K. Jalali, Ph.D., FRES Principal Scientist NBAIR, Bangalore
External
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Preamble
Ways- intrinsic competition
Mechanisms in intrinsic competition
Case studies
Solitary Vs. Gregarious intrinsic competition
Fitness of progeny in intrinsic competition
PART I
PART II
PART III
INDEX
Conclusion References
Competition Strategies in intrinsic competition
Factors effecting intrinsic competition
Discussion
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PART I
PREAMBLE
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Integrated Pest Management Legal control Physical control Cultural control Mechanical control “Biological control” Chemical control Host plant resistance Biological control : “Parasitoids”
Predators Pathogens
Cassava mealybug-Africa Papaya mealybug-India Cottony cushiony scale- California
Parasitoid
Immature stage is parasitic Adult is free living
“Insect parasite of an arthropodan “
Adult
“Obligatory but facultative”
-Jervis, 2005
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Host/prey
Phytophagy
Parasitoid/Predator
Patch
Clutch size
Phytophagous pest Parasitoid/PredatorX-Jervis, 2005
Host
Herbivore/ Phytophagy
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PART II
Ways- intrinsic competition
Mechanisms in intrinsic competitionSolitary Vs. Gregarious intrinsic competition
Fitness of progeny in intrinsic competition
Competition Strategies in intrinsic competition
Factors effecting intrinsic competition
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"Competition is a manifestation of the struggle for existence in which two or more organisms of the same or of different species exert a disadvantageous influence upon each other because their more or less active demands exceed the immediate supply of their common resources." Predators- Broad host range Parasitoids- Narrow host range
1. Host habitat finding 2. Host finding 3. Host acceptance 4. Host suitability
Host/Patch marking
Requisite host in less Volatility of the marking pheromone Sharing of the hosts in a patch Pro ovigeny condition
Failure of host discrimination
“Competition”
Despite of having egg, larval, egg larval parasitoids Despite of having egg, larval, egg larval parasitoids
Plant-herbivore-parasitoid systems Integration of the parasitoids-successful pest control -Paul Debach, 1973
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Two types of competition: 1. Extrinsic competition 2. Intrinsic competition
Extrinsic competition Intrinsic competition
Among free-living adults Among immature parasitoids
Life cycle of a parasitoid
Egg to egg - xEgg to I instar larvae I instar to late instar
Harvey et al., 2013
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Kill the host Static resources-quality
Many individuals
Single individual
Outside of the host Perforate the cuticle Imbibe the fluids Large with yolk eggs
Allow it grow the host Resources dynamic
Tissue, Haemolymph Direct contact Yolkless eggs-extra membrane Total feeder/haemolymph/fat
-Yamamoto et al.,2007 and Harvey et al., 2013
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Microplitis mediator
Solitary larval endoparasitoid Gregarious larval endoparasitoid
Cotesia glomerata
Diadegma semiclausum
Solitary larval endoparasitoid
-Gols and Harvey, 2009
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Two ways 1. Superparasitism 2. Multiparasitism
Superparasitism: Parasitism of a host by parasitoids of the same speciesMultiparasitism:Parasitism of a host by parasitoids of the different species
Antagonistic interactions - monopolize host resources.Parasitoids do not hesitate to oviposit in already parasitized hosts Self and Conspecific Heterospecific Superparasitism delays progeny
development of immature parasitoids Smaller parasitoid progeny Competitive displacement
-Bai and Mackauer, 1992
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Two mechanisms 1. Physical defense 2. Physiological suppression
On hatching - move actively-abdomen and tail
Embeds its mandibles into the body of the other
Leakage of body fluid from the punctured cuticle(Whole of the bitten larva becomes considerably shrunken).
Larva-susceptible to the blood reactions of the host
Encapsulation by haemocytes, deposition of melanin
Distance between the members of each pair of spots distance between the mandibles
Aggregation of host phagocytes around the wound
6 days this enlarges considerably and either encloses the whole larva in a loose capsule or forms a tubular capsule around itHarvey et al., 2013
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Embedding mandibles Melanin-points of wounding partial encapsulation
Melanization and a tubular haemocyte capsule
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Cessation of feeding, shrinking and stunting of the larvae Encapsulation (Sub spherical/Regular outline compact capsule) Improper eclosion of the larvae
No mandible attack only contact through the haemolymph
What ?
How ?
Secretion of the toxic substance Stimulation host blood to produce more phagocytes Conditioning-haemolymph-Excretory, salivary secretions
Encapsulation
Larvae Adult I. Larvae
Harvey et al., 2013
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DIFFERENTIATION ?
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II. Adult Molecular tug-of-warTeratocytes, Venoms and Polydnaviruses (PDVs)
Passive and Active
Egg coating
Hatching Regulation of host
growth Synergistic effect Trophic function
-Asgari, 2006 and Harvey et al., 2013
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Venom with calyx fluid Pupal stage 10-100 kda 6 major proteins
Ectoparasitoids (Venom) Endoparasitoids (Venom+PDVs)
Paralysis, Morphology, viability, and immune function of haemocytes Host immune suppression Haemocytes spreading
Indirect effect of venom Promoting PDV infection Synergistic effect
-Asgari, 2006 and Harvey et al., 2013
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His research on the underlying mechanisms of insect immunity culminated in the treatise, The Cellular Defence Reactions of Insects
Placing washed eggs of the parasitoid Venturia (Nemeritis) canescens into living larvae of its host Ephestia kuehniella
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Calyx proteins
Ichnovirus Bracovirus
Venturia (Nemeritis) canescensGenome-integrated wasp symbiontsSlavicek, 2012
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Slavicek, 2012
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Slavicek, 2012
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Braco Ichno
Slavicek, 2012
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Phenoloxidase enzymes convert tyrosine to dopa, and dopa to dopaquinone, leading to the formation of melanin;
Slavicek, 2012
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1.Time of oviposition2.Environmental temperature 3.Host species 4.Host feed-Nasty host hypothesis
Time of ovipositionSelf/Conspecific/Hetero
Host species-The caterpillar Host feed-caterpillar feeds
-Fisher, 1961; Weil and Sagarzazu 1998 and Harvey et al., 2005
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I. Oviposition at the same time Horogenes chrysostictos Nemeritis canescens
Ephestia sericarium
Methodology 15 healthy mature larvae for 4 hr
6 replicates @ 25 0C (90 Larvae)
4 DAO-dissections-Observations
-Fisher, 1961
41 of the Ephestia containing both species 36-only one larva 24 –Horogenes; 12-Nemertis
Emergence time :Horogenes : 59 + 2 hr; Nemertis : 69 + 2 hr
Changed the experimentInjection method
6 Days after injection observations
8 hosts unsuccessful16 died before conclusion of the exp. 23/18
Ratio: 1: 1 Active & undamaged Bitten Melanin deposition
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ii) Oviposition at different times
-Fisher, 1961
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-Fisher, 1961
The effect of temperature on development of the eggs and I-instar larvaeEgg
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Larva
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Endoparasitoids ecological niches-host insects Ephestia sericarium Ephestia elutella Ephestia cautella “Galleria mellonella” Achroia grisella
Horogenes chrysostictos Nemeritis canescens Development is slower
11 days- I instar moult ( 6 days in Ephestia) 32-33 days delayed emergence (22 days ) Horogenes did not effect the development
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Herbivore
Parasitoid foraging
“HIPVs”Host may seek different food plants Quality of food plants is important Nutrients and the toxins - Performance of the host
“Direct indirect effects”
Two types of interactions
Plant-Herbivore-Parasitoid Parasitoid-Herbivore-Plant
koinobiont endoparasitoids Network of interactions Asymmetric fashion
Phytotoxins adaptations - influence intrinsic comp.Stronger competitor in preferred than additional. Compramisation of immune response
- Gols and Harvey 2009 and Harvey et al., 2013
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“Safe host/heaven hypothesis”
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Solitary parasitoids Strong mandibles in the first instar larvae. Well developed caudal appendage/tail
Without any mandibles Lack of caudal appendage/tail Larvae occupy much more confined spaces Without necessarily antagonistic encounters Scramble competition
Gregarious parasitoids
Par
asito
id lo
ad
Parasitoid fitness
- Harvey et al., 2013
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Antagonistic interactions Winning competitor-Elimination of superneumeries
Effect of fitness Survival – Superparasitism/Multiparasitism/Simple Adult size-Decreased Developmental time increased Adult longevity decreased
Idiobionts-Per capita resources are lessKoinobionts-Regulate the host feeding rate and growth
-Strand et al., 1988 and Harvey, 2005 and Harvey et al., 2013
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PART III Case studies
Conclusion ReferencesDiscussion
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1
Microplitis croceipus Cardiochilus nigriceps Campoletis sonorensis Chelonus insularis
“Multi-interspecific-endoparasitic competition”
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Methodology
Tobacco budworm
Glass container
Paper towel
Cut squares (20-30 eggs)
3/4 females4 hours
Long vial Artificial diet
Petri dish
Larvae Second stage larvae
Expose to the individual larval endoparasitoid
Emergence
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Emergence of larval parasitoids
Results and Discussion Fate of larval endoparasitoids
Host discrimination M. Croceipus; C. sonorensis Physical combat No advantage
Larger size Mobility Poor mandibles
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Cardiochilus is a poor competitor
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2
Cotesia flavipes Cotesia sesamiae “Multi-Intra/interspecific-endoparasitic competition” Chilo partellus
Sesamia calamistis
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3
“Multi-Intra/interspecific-endoparasitic competition”
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Methodology
Collection-Mythimna separata and parasitoids
Artificial diet, INSECTA-LFS-16:8 hrs. Photoperiod, 25 ±1 o C
Mating and maturation-oviposition activity @ room temp., 30 % sugar solution
100 M. separata larvae parasitisation individually
Reared in the plastic boxes
Dissection-HEPES buffered saline solution
Observations: oviposition to cocoon formation
Morphological observations-SEM
Pair wise contest (1, 24, 48 and 96 hrs intervals
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Important component is bio control: parasitoids, predators, pathogens Parasitoids important to study to understand the parasitoid-herbivore-
plant interaction Trends like development of insecticide, thermo tolerant, Diapause,
Genetic engineering & Competition Host requisite in less, provigeny condition- failure of host
discrimination Two types: extrinsic and intrinsic Extrinsic: free living adults and intrinsic: immature stages Idiobionts, Koinobionts, Ecto, Endo, Solitary and Gregarious Competition: between the egg and I instar and I instar and later instars Super and multi parasitism –Ways of competition Mechanisms: physical and physiological Physical by the mandibles and physiological by larva and adult Larvae: toxic substances, stimulation and conditioning of the
haemolymph Adult: Teratocytes, Venom, PDVs
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Factors: Time of Oviposition, Temperature, Host and host feed (Nasty and Safe heaven hypothesis)
Based on the competition the fitness of the progeny would vary. Winning parasitoid fitness lesser than the simple parasitic progeny Optimum load crossed the scramble competition takes place in the
gregarious parasitoids In solitary parasitoids I instars use mainly the sickle shaped mandibles
for defense In gregarious parasitoids mandibles not well developed and dependent
on the physiological suppression Later instars also dependent on the physiological suppression
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“As the parasitoids are involved in the competition because of their narrow host range. Biocontrol programmes should concentrate on the best integration methods involved with egg, larval and pupal parasitoids and their time of release. Molecular techniques should develop to identify the multiparasitic hosts”.
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A VIDEO ON INTRINSIC COMPETITION
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Vinson, S. B., J. R. Ables. 1980. Interspecific competition among endoparasitoids of tobacco budworm larvae (Lepidoptera: Noctuidae). Entomophaga. 25(4): 357-362.
Harvey, J.A., E.H. Poelman and T. Tanaka. 2013. Intrinsic inter and intraspecific competition in parasitoid Wasps. Annu. Rev. Entomol. 58: 333-351.
Sallam, M.N., A. William, Overholt and E. Kairu. 2002. Intraspecific and interspecific competition between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), gregarious larval endoparasitoids of lepidopteran stem borers. Bioc. Sci. and Tech. 12: 493-506.
Magdaraog, P.M., J.A. Harvey, T. Tanaka and R. Gols. 2012. Intrinsic competition among solitary and gregarious endoparasitoid wasps and the phenomenon of resource sharing. Ecological Entomology, 37: 65-74.
Gols, R. and J.A. Harvey.2009. Plant-mediated effects in the Brassicaceae on the performance and behaviour of parasitoids. Phytochem Rev., 8:187-206.
Fisher. R.C. 1961. A study in insect multiparasitism II. The mechanism and control of competition for possession of the host J. Exp. Biol., 38: 605-628.
Asgari, S. 2006. Venom proteins from polydnavirus-producing endoparasitoids: Their role in host-parasite interactions. Archives of Insect Biochemistry and Physiology, 61:146-156.
Mayhew, P.J. and J.J.M. Van Alphen.1999. Gregarious development in alysiine parasitoids evolved through a reduction in larval aggression. Animal Behaviour, 58: 131-141.
Harvey, J.A., F. Pashalidou, R.Soler and T. Martijn Bezemer.2011. Intrinsic competition between two secondary hyperparasitoids results in temporal trophic switch, Oikos, 120: 226-233.
EC. Lampert, LA. Dyer and M. D. Bowers. 2010. Caterpillar Chemical Defense and Parasitoid Success: Cotesia congregata parasitism of Ceratomia catalpa. J. Chem. Ecol., 36: 992-998.
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What is the distance between two mandibles if the distance between the two melanin spots on the body of that larva is 250 µm ? A)250 µmB)500 µmC)1200 µmD)1500 µm
Answer ?