influence of plant age, soil moisture, and temperature ... · influence of plant age, soil...
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Influence of Plant Age, Soil Moisture, and
Temperature Cycling Date on
Container-Grown Herbaceous Perennials
Sarah L. Kingsley-Richards
Master‟s Candidate, Plant & Soil Science
University of Vermont
March 16, 2011
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Perennials Woody Plantsmaintain meristem potential true dormancy*
stimulated to acclimate by
low temperatures
stimulated to acclimate by
shortening day length
photosynthesis during the early
stages of acclimation
no photosynthesis once
acclimation begins
Stage #1 above freezing
Stage #2 below freezing
Stage #1 cease growth
Stage #2 low temperatures
Stage #3 temperatures < -30ºC
*Dormancy = the temporary suspension of visible growth of
any plant structure containing a meristem
vs.
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Plant Cell Structure
• Cell wall*
• Cell membrane
• Cytoskeleton
• Organelles
• nucleus
• chloroplasts*
• mitochondria
• etc.
*unique to plants
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Acclimation Process
1. Membrane lipids shift to a solid state
2. Cytoskeleton is rearranged
3. Membrane proteins cease functioning Ion leakage
4. Chain of enzyme reactions and expression of genes
5. Abscisic acid accumulation
6. Photosynthesis continues
7. Cell solutions become concentrated, particularly with sugars
remain liquid below the freezing point of water = supercooling
As long as the temperature drops slowly (1-2ºC/hour),
this state will normally persist in hardy plants.
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Rapid Temperature Drop = Ice in cell
almost always fatal to cell
• unless very small ice crystals
• broken cell membrane leads to cell death
• cells may synthesize „anti-freeze‟ proteins that
regulate the formation and size of ice crystals
Ice formation outside of the cell is common
• lower solute concentration and more nucleating sites
• most hardy plants tolerate it
• possible to „dry up and die‟
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Plant Survival Factors
• genetics/species variation
• environmental cues
• rate at which temperatures drop
• lowest temperature and how long
• temperature cycling
• insulation
• plant age
• plant health
• moisture content
• fertility
• pot size
Plants in pots have less tolerance
of extreme-temperatures
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Why Freeze Plants in Pots?
• Nursery growers often want to over-winter plants
• Worst possible conditions for survival
• Easy to control freezing conditions
• Assess whole-plant performance
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The Hardiest Perennial?
P. Becquerel, 1932, Ranunculus tubers
18 days in liquid nitrogen at -190ºC (-310ºF)
• 9% moisture survived
• 30-50% moisture died
The lowest temperature recorded on the planet
-91ºC (-132ºF) in Antarctica
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Three sets of experimentsTo assess cold hardiness of container-grown herbaceous perennials
under the following influences:
*data not presented –see thesis Appendix
2004-2005 2005-2006 2006-2007
Plant Age X X
Soil Moisture X X
Temperature
Cycling DateX X
Freeze Only* X X X
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Plant Age
Geranium x cantabrigiense 'Karmina' (only in 2005-2006)
Leucanthemum x suberbum 'Becky'
Sedum 'Matrona'
-NEW plants were obtained as liners at the beginning of the growing
season
- OLD plants had been established in pots for one prior growing season
and were a year old at the beginning of the growing season
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Soil Moisture
2004-2005
Coreopsis 'Tequila Sunrise'
Geranium x cantabrigiense 'Cambridge'
Heuchera 'Plum Pudding'
2006-2007
Carex oshimensis 'Evergold'
Carex morrowii 'Ice Dance'
Carex laxiculmis 'Hobb' (Bunny Blue™)
Volumetric water content (VWC) of potting media
was monitored over the growing season.
- WET plants maintained above 10% VWC
- DRY plants maintained below 10% VWC
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Temperature Cycling
Geranium x cantabrigiense 'Cambridge' (only in 2004-2005)
Leucanthemum x suberbum 'Becky'
Sedum 'Matrona' (only in 2005-2006)
In NOVEMBER, JANUARY, and MARCH
2-CYCLE = -3C 24h, +3C 24h, -3C 24h, +3C 24h
1-CYCLE = -3C 24h, +3C 24h
0-CYCLE = no temperature cycling
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Establishing Plants
• Thirty plants were established for each species for each factor
either from liner plugs or previously established plants.
• Plants were allowed to establish over a normal growing season
in the UVM greenhouse under ambient temperature.
• Plants were watered as needed throughout study (with the
exception of the moisture study prior to freezing).
• Water soluble fertilizer was applied weekly throughout growing
season (Jack‟s Professional 17-14-17 & STEM soluble trace
elements).
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
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Acclimating Plants
• Temperatures in greenhouse were reduced beginning in
October of each year at a rate of 3°C (5°F) per week until
temperatures of 3-5°C (37-41°F) were reached at the end of
November.
• This low temperature was maintained in the
greenhouse until spring when the temperature
was increased by the same increments
beginning in April of each year until
ambient temperature was reached.
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Freezing Plants
• During the month of January, the
thirty plants were randomly divided
into five, six-pot replicates per factor.
• Plants were pruned back to 1” of pot
edge in most cases prior to freezing.
• Controlled freezing of each set of
replicates to temperatures of
-2, -5, -8, -11, and -14ºC was
performed in chest freezers.
(28, 23, 18, 12, and 7ºF)
Freezing for the temperature cycle study was also performed
in November and March.
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Plant Age
NEW
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
OLD
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
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Soil Moisture
WET
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
DRY
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
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Temperature
CyclingN
O
V
E
M
B
E
R
0-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
1-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
2-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
J
A
N
U
A
R
Y
0-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
1-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
2-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
M
A
R
C
H
0-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
1-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
2-cycle
-2ºC -5ºC -8ºC -11ºC -14ºC
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
X X X
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Basic Freezing Protocol
1. Load all plants into freezer
with -14°C on bottom to -2°C on top.
2. Set freezer to -2°C for 48 hours.
3. Remove -2°C plants, set freezer to -5°C for 2.5 hours.
4. Remove -5°C plants, set freezer to -8°C for 2.5 hours.
5. Remove -8°C plants, set freezer to -11°C for 2.5 hours.
6. Remove -11°C plants, set freezer to -14°C for 2.5 hours.
7. Remove -14°C plants.
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Cycling Freezing Protocol
1. Put the 2-cycle plants in freezer at -3°C for 24 hours.
2. Increase freezer temperature to +3°C for 24 hours.
3. Add the 1-cycle plants to freezer and set to -3°C for 24 hours.
4. Increase freezer temperature to +3°C for 24 hours.
5. Sort all plants, including 0-cycle, by target freezing temperature.
6. Follow steps of Basic Freezing Protocol
using -3°C instead of -2°C.
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Surviving Plants
• Plants returned to the greenhouse
were allowed to re-grow.
• Visual rating of plants was
performed in June/July on a scale of
1-5, with 1 being dead and 3-5 being
increasingly salable plants.
• Dry weight of each plant was
measured by cutting back all new
growth to 1” of pot edge, bagging in
paper bags, drying in drying room at
HRC, then measuring the bag.
• Data from rating and dry weight
analyzed using SAS.
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Plant Age Results
• Geranium x cantabrigiense 'Karmina'
• no effect on either rating or dry weight
• Sedum 'Matrona'
• no effect on dry weight
• Ratings higher for older plants than younger plants in the
first year and higher for younger plants than older plants in
the second year
• Leucanthemum x superbum 'Becky'
• rating and dry weight both generally higher for younger
plants than older plants
In general, when an effect was observed, younger plants
produced more growth and rated higher in quality
following exposure to freezing temperatures.
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Ratinga
2005-2006 2006-2007
Treatment
temp °C
Younger
plants b
Older
plants c Mean (SEM
d)
Younger
plants
Older
plants Mean (SEM)
-2 3.00 2.50 2.75 (0.22) a e 4.33 Aa
k 1.50 Bc 2.92 (0.45)
-5 3.17 2.33 2.75 (0.22) a 3.83 ab 3.67 a 3.75 (0.13)
-8 2.50 1.67 2.08 (0.26) ab 3.67 Ab 3.00 Bb 3.33 (0.22)
-11 1.67 1.50 1.58 (0.15) bc 4.00 Aab 3.00 Bb 3.50 (0.15)
-14 1.00 1.33 1.17 (0.11) c 2.00 c 1.67 c 1.83 (0.11)
Mean
(SEM)
2.27 A f
(0.19)
1.87 B
(0.14)
3.57
(0.16)
2.57
(0.18)
ANOVA significance:
df g F
h p-value
i df F p-value
Temperature 4 14.35 <.0001 4 29.18 <.0001
Plant age 1 5.81 0.0197 1 64.29 <.0001
Interaction 4 1.77 0.1488 4 14.82 <.0001
Effect of plant age and freezing temperatures on quality RATING of
Leucanthemum x superbum 'Becky' regrowth .
Younger plants rated
higher in quality
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Effect of plant age and freezing temperatures on DRY WEIGHT of
Leucanthemum x superbum 'Becky' regrowth .Dry weight (g)
2005-2006 2006-2007
Treatment
temp °C
Younger
plants b
Older
plants c Mean (SEM
d)
Younger
plants
Older
plants Mean (SEM)
-2 1.61 0.75 1.18 (0.22) a e 3.67 Aa
k 0.62 Bb 2.14 (0.50)
-5 1.50 0.78 1.14 (0.14) a 3.32 Aa 1.36 Ba 2.34 (0.32)
-8 0.87 0.24 0.56 (0.19) b 2.53 Ab 1.01 Bab 1.77 (0.26)
-11 0.29 0.09 0.19 (0.07) bc 3.10 Aab 0.94 Bab 2.02 (0.35)
-14 0.00 0.07 0.03 (0.02) c 1.23 Ac 0.35 Bb 0.79 (0.18)
Mean
(SEM)
0.86 A f
(0.15)
0.39 B
(0.07)
2.77
(0.18)
0.86
(0.10)
ANOVA significance:
df g F
h p-value
i df F p-value
Temperature 4 17.95 <.0001 4 16.25 <.0001
Plant age 1 17.84 0.0001 1 201.10 <.0001
Interaction 4 2.46 0.0571 4 7.11 0.0001
Younger plants had
higher dry weight
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Soil Moisture Results
• Coreopsis 'Tequila Sunrise' & Carex morrowii 'Ice Dance'
• no effect on either rating or dry weight
• Geranium x cantabrigiense 'Cambridge' “wet”
& Heuchera 'Plum Pudding' “dry”
• no effect on dry weight but ratings were higher
• Carex laxiculmis 'Hobb' (Bunny Blue™)
• dry weight was higher for “dry” plants
• Carex oshimensis 'Evergold'
• effects were inconclusive
A general effect of soil moisture level on survival and
salable quality following exposure to freezing
temperatures was not apparent in these studies.
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Temperature Cycling Results -Cycles
• Geranium x cantabrigiense 'Cambridge'
• advantageous effect in November
• Sedum 'Matrona'
• advantageous effect in March
• Leucanthemum x superbum 'Becky'
• advantageous effect in November in the first year
& in all months in the second year
A general tendency was observed for cycles to increase
hardiness
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Temperature Cycling Results - Date
• Geranium x cantabrigiense 'Cambridge'
& Sedum 'Matrona'
• more tolerant of cycling and freezing in January
• Leucanthemum x superbum 'Becky'
• more tolerant of cycling in January in the second year
A general tendency was observed for hardiness to be highest
in January than November or March
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Table 4.1. Effect of temperature cycling date and freezing temperatures on regrowth ratinga of Geranium x catabrigiense 'Cambridge'.
November 2004-2005 January 2004-2005 March 2004-2005
Treatment
temp °C 0-cycles b
1-cycle c 2-cycle
d Mean (SEM
e) 0-cycles 1-cycle
2-cycle
Mean (SEM) 0-cycles 1-cycle
2-cycle
Mean (SEM)
-2 3.00 2.83 2.83 2.89 (0.18) a f 2.83 2.50 3.17 2.83 (0.20)
3.17 2.67 2.17 2.67 (0.16) a
-5 2.50 2.67 2.83 2.67 (0.14) a 2.67 2.50 2.83 2.67 (0.14) 2.67 2.50 2.17 2.44 (0.15) ab
-8 2.67 3.00 3.00 2.89 (0.16) a 2.83 2.00 2.33 2.39 (0.14) 2.33 2.67 2.33 2.44 (0.12) ab
-11 2.17 3.00 2.67 2.61 (0.14) ab 3.00 2.83 2.67 2.83 (0.15) 2.17 2.17 2.17 2.17 (0.12) ab
-14 1.33 2.50 2.17 2.00 (0.18) b 3.00 2.50 2.83 2.78 (0.17) 1.83 2.33 1.67 1.94 (0.13) b
Mean
(SEM)
2.33 B g
(0.16)
2.80 A
(0.13)
2.70 AB
(0.10)
2.87
(0.15)
2.47
(0.10)
2.77
(0.11)
2.43 AB
(0.12)
2.47 A
(0.09)
2.10 B
(0.11)
ANOVA significance:
df h F
i p-value
k df F p-value df F p-value
Temperature 4 5.51 0.0006 4 1.33 0.2680 4 4.69 0.0020
Month 2 4.18 0.0190 2 2.75 0.0706 2 4.08 0.0208
Interaction 8 1.07 0.3929 8 0.60 0.7713 8 1.32 0.2451
aRating scale 1 = Dead, no regrowth, 2 = No flowering stems and minimal regrowth, 3 = 0-2 flowering stems and regrowth extending over edge of pot, 4 = 3-5 flowering stems and
regrowth equal to or greater than above, 5 = 6 or more flowering stems and regrowth as above. b0 cycle plants were not subjected to temperature fluctuations prior to controlled freezing at lower temperatures.
c1 cycle plants were held at -2C for 24 hours, then held at 3C for 24 hours prior to controlled freezing t lower temperatures.
d2 cycle plants were held at -2C for 24 hours, then held at 3C for 24 hours, then held at -2C for 24 hours, then held at 3C for 24 hours prior to controlled freezing at lower temperatures.
eSEM=Standard Error of the Mean.
fTreatment temperature means with a lowercase letter in common are not significantly different according to Tukey‟s procedure (p=0.05).
gWhere no interaction present between factors, month means with a capital letter in common are not significantly different according to Tukey‟s procedure (p=0.05).
hdf=degrees of freedom.
iF=F distribution.
kp-value=level of significance.
Effect of temperature cycling date and freezing temperatures on
quality RATING of Geranium x cantabrigiense 'Cambridge' regrowth
-3
Cycles have no
effect in January
= more hardy
Cycles increase
hardiness in
November
(more than in March)
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Table 4.2. Effect of temperature cycling date and freezing temperatures on regrowth dry weight of Geranium x catabrigiense 'Cambridge'.
November 2004-2005 January 2004-2005 March 2004-2005
Treatment
temp °C 0-cycles b
1-cycle c 2-cycle
d Mean (SEM
e) 0-cycles 1-cycle
2-cycle
Mean (SEM) 0-cycles 1-cycle
2-cycle
Mean (SEM)
-2 0.27 0.38 0.40 0.35 (0.03) a f 0.42 0.35 0.22 0.33 (0.03) 0.40 0.46 0.25 0.37 (0.03) a
-5 0.19 0.28 0.28 0.25 (0.03) ab 0.27 0.34 0.30 0.30 (0.03) 0.32 0.32 0.26 0.30 (0.02) ab
-8 0.22 0.26 0.34 0.27 (0.03) a 0.27 0.18 0.30 0.25 (0.04) 0.20 0.29 0.31 0.26 (0.03) b
-11 0.12 0.33 0.34 0.26 (0.03) a 0.30 0.26 0.27 0.28 (0.03) 0.28 0.32 0.26 0.29 (0.03) ab
-14 0.6 0.16 0.22 0.15 (0.02) b 0.34 0.33 0.30 0.32 (0.03) 0.17 0.32 0.17 0.22 (0.03) b
Mean
(SEM)
0.17 B g
(0.02)
0.28 A
(0.02)
0.31 A
(0.02)
0.32
(0.03)
0.29
(0.02)
0.28
(0.02)
0.27 AB
(0.02)
0.34 A
(0.02)
0.25 B
(0.02)
ANOVA significance:
df h F
i p-value
k df F p-value df F p-value
Temperature 4 6.81 <.0001 4 1.05 0.3862 4 4.56 0.0024
Month 2 12.27 <.0001 2 0.65 0.5226 2 5.59 0.0054
Interaction 8 0.54 0.8200 8 1.14 0.3496 8 1.67 0.1195
b0 cycle plants were not subjected to temperature fluctuations prior to controlled freezing at lower temperatures.
c1 cycle plants were held at -2C for 24 hours, then held at 3C for 24 hours prior to controlled freezing t lower temperatures.
d2 cycle plants were held at -2C for 24 hours, then held at 3C for 24 hours, then held at -2C for 24 hours, then held at 3C for 24 hours prior to controlled freezing at lower temperatures.
eSEM=Standard Error of the Mean.
fTreatment temperature means with a lowercase letter in common are not significantly different according to Tukey‟s procedure (p=0.05).
gWhere no interaction present between factors, month means with a capital letter in common are not significantly different according to Tukey‟s procedure (p=0.05).
hdf=degrees of freedom.
iF=F distribution.
kp-value=level of significance.
Effect of temperature cycling date and freezing temperatures on
DRY WEIGHT of Geranium x cantabrigiense 'Cambridge' regrowth
Cycles have no
effect in January
= more hardy
Cycles increase
hardiness in
November
(more than in March)
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In Summary
• Younger plants produced more growth and rated higher
• A general soil moisture effect was not apparent
• Cycles can increase hardiness
• Hardiness highest in January
NOTE: These conclusions are generalized.
Individual cultivars may react differently
under other cultural conditions.
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Acknowledgements
• Leonard Perry
• Scott Costa
• Kathleen Liang
• PSS faculty and staff
.
• Aaron Richards
• Family & friends
Thank You!
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