Genecology and Seed Zones for Bluebunch
Wheatgrass and Prairie Junegrass
Brad St.Clair
Francis Kilkenny
USDA FS Pacific Northwest Research Station, Corvallis, OR
Photo: Berta Youtie
2012 Great Basin Native Plant Selection and Increase Project Annual Meeting
Collaborators
R.C. Johnson
USDA ARS Western Regional Plant Introduction Station, Pullman, WA
Nancy Shaw
USDA FS Rocky Mountain Research Station, Boise, ID
Matt Horning
USDA FS Region 6, Bend, OR
Dale Darris
USDA NRCS Plant Materials Center, Corvallis, OR
Many others who provided seed from many populations
Photo: Berta Youtie
Adaptation
The evolutionary process whereby an population
becomes better suited to its environment
A feature which is especially important for an
organism’s survival and reproduction; a product of
natural selection in a given environment (adaptive
trait)
1.
2.
•
•
•
Heritable
Functional
A result of natural selection
Adaptedness = the state of being adapted; the degree
to which an organism is able to survive, grow, and
reproduce in a given environment (related to fitness)
Evidence for adaptation comes from:
1.
2.
3.
4.
Correlation between a character and environmental
factors - the same form occurs in similar environments
(genecology studies)
Comparisons of naturally-occurring variants in
environments where they are hypothesized to function
as adaptations (reciprocal transplant studies)
Direct evidence from altering a character to see how it
affects function in a given environment
In population genetics, deviation from what would be
expected from neutrality (e.g., Fst outliers, Tajima’s D)
Adapted from West-Eberhard 1992, Endler 1986
Common Garden Studies
Common-garden experiments are used to separate
genetic from environmental effects
Environment is more uniform
• “E” is smaller
• Phenotype (P) more closely reflects the genotype (G)
P=G+E
Genecology Studies
Genecology
• The study of intraspecific genetic variation of
plants in relation to source environments
(Turresson 1923)
• Seeks correlations between “plant type” and
“habitat type”
• Consistent correlations are taken to indicate
adaptation as determined by natural selection
Example: Douglas-fir Genecology Study
Grow families in a
common environment
Measure many
adaptive traits
Collect
seed
from
many
trees
GIS
Combination of Variables, Primarily Growth
Douglas-Fir of Western OR and WA
3
2
1
0
Traits vs
source
environment
-1
-2
-3
-4
-5
-10
-8
-6
-4
-2
0
2
December Minimum Temperature
4
6
Advantages/disadvantages of genecology studies
•
•
•
•
May be done in a short timeframe
Greatly minimizes environmental variation
Can sample many source locations from a wide range of environments
Can produce maps of adaptive traits/climates for easy visualization
and manipulation
But,
• Assumes we have measured all the most important
adaptive traits
• May be difficult to synthesize results of many traits
• Comparisons between sites or years assume local is
best (risk of maladaptation, LSD)
• Not a direct test of adaptation
Reciprocal Transplant Studies
Populations from a range of source environments are
evaluated in the same or similar range of test
environments
Can generate models to predict adaptation as a function of:
1.
2.
3.
4.
planting environments (response function)
source environments (genecology functions)
difference between a planting and source environments
(transfer functions)
both the planting environment and source environment together
(universal response functions)
Mather
El. 1,400 m
Potentilla glandulosa from three
different elevations planted at
three different elevations
(Clausen, Keck & Hiesey 1940)
Stanford
El. 35 m
Native to
Timberline
El. 3,030 m
Classical studies of Clausen, Keck, & Hiesey
Stanford
El. 35 m
Mather
Timberline
El. 1,400 m El. 3,030 m
Grown at
Pinus contorta
provenance trial
140 provenances sampled
60 test sites
Variation among
four populations
at one test site
Manning
49 N
Albreda
52 N
Champion
49 N
Takhini
60 N
11
Response functions derived from lodgepole pine
provenance tests in British Columbia
from Wang et al. 2006. Use of response functions in
selecting lodgepole pine populations for future climate.
Global Change Biology 12: 2404-2416.
Transfer functions
Fig. 7 Individual transfer functions (the trend curves) for six test sites from a range of mean annual
temperatures (MATs). The vertical dashed line indicates the site MAT.
Universal Response Function
Response function:
Yi = b0 + b1X1i + b2X21i + ei
Genecology/transfer function:
Yj= b0 + b1X2j + b2X22j + ej
Combined:
Yj= b0 + b1X1i + b2X21i + b3X2j + b4X22j + b5X1j X2j + eij
Add geographic effects:
Yj= b0 + b1X1i + b2X21i + b3X2j + b4X22j + b5X1j X2j +
b6X3i + b7X23j + eij
Wang, O’Neill and Aitken. 2010.
Ecological Applications
Advantages/disadvantages of reciprocal
transplant studies
• Direct test of adaptation
• Can test hypothesis of local adaptation
• Can better model variables of direct interest such as stand
productivity
But,
• Takes a long time for results
• Expensive to test over many planting sites and seed sources
• Few samples makes it difficult to interpolate between locations to
adequately model response or transfer functions, or draw maps
• Does not provide information on components of adaptation and,
thus, traits that might be useful for selection of populations or
individuals
Objectives
1.
Explore genetic variation among populations
from a wide range of source environments in
the inland West
2. Relate genetic variation to environmental
variation at source locations
3. Develop seed transfer guidelines
4. Compare native variation to common cultivars
Goal: adapted, diverse plant populations
for restoration
Bluebunch wheatgrass (Pseudoroegneria
spicata): Population Sampling
Collections from:
114 populations
2 families per population
5 cultivars
Hanford Reserve, Central Washington
Bluebunch wheatgrass:Test Sites
Planted at three common garden test sites in 2006:
Central Ferry, WA – warm, dry
Lucky Peak Nursery, ID – cooler, dry
Pullman, WA – cold, wet
6 replications, 1 plant per family per rep
4,752 total plants
Transplanting Bluebunch Wheatgrass,
Lucky Peak, Fall 2006
Bluebunch wheatgrass: Measurements
Eighteen traits measured in 2007 & 2008:
• Size – biomass, crown width, height, regrowth biomass
• Fecundity – inflorescence number
• Phenology – heading, anthesis, seed maturation, germination
• Morphology – plant form, leaf width, leaf length, leaf color,
leaf pubescence, culm length, spike length, spikelet number,
awn length
Procedure
1.
2.
3.
4.
5.
6.
7.
Sample populations from many diverse environments
Measure traits in a common environment
ANOVA to explore sources of variation and
interactions; particularly interested in traits with high
population variation
Principal component analysis to reduce number of traits
Correlations and regressions to explore relation of
traits to climate
Use GIS and regressions to produce maps of genetic
variation
Overlap PC maps to delineate areas of similar
populations = seed zones
Bluebunch: ANOVA
Trait
Pop
Site
Year
Pop*site
Pop*year
Site*year
DRYWT
15.51**
80.7**
2690.48**
5.59**
9.51**
939.13**
CW
HT
REGRWT
REGRHT
15.05**
6.95**
11.58**
6.8**
89.62**
94.09**
39.72**
72.18**
8923.33**
1.98
2876.2**
0
2.75**
2.37**
2.35**
3.16**
5.34**
1.94**
9.72**
1.36**
1003.37**
71.47**
969.53**
105.64**
INFLNO
8.73**
119.48** 4044.67**
3.45**
6.6**
2087.96**
CULMLNG
SPKLNG
INFLLNG
SPKNO
LFLNG
LFWD
LFRATIO
LFCOL
6.18**
0.82
1.89**
8.78**
6.05**
3.34**
12.22**
4.97**
1031.27**
27.05**
391.18**
266.57**
73.25**
0.56
14.16**
1.23
191.42**
10.92**
0.35
372.33**
338.85**
259.51**
1.27
64.8**
5.09**
0.91
1.19*
1.54**
1.86**
0.68
1.77**
1.21*
2.07**
1.06
1.17
1.96**
2.11**
0.67
1.55**
1.36**
135.35**
1.49
4.6*
70.81**
481.89**
126.41**
71.12**
13.11**
LFPUB
HABIT
AWNS
10.29**
5.27**
18.05**
0.03
12.63**
0.21
1710.47**
1.27
17.69**
3.16**
2.39**
1.43**
1.78**
2.53**
1.21
293.52**
143.55**
1.05
HEAD
6.28**
784.14** 1805.41**
1.75**
2.05**
424.14**
BLOOM
MATURE
GERM3
7.23**
2.66**
2.82**
1615.44** 3232**
376.15** 138.68**
34.6**
-
1.3**
1.39**
1.41**
1.85**
1.98**
-
780.82**
217.15**
-
Bluebunch: Percent trait variation due to
population of origin
SITE=CF
year=2007
DRYWT
CW
HT
REGRWT
REGRHT
INFLNO
CULMLNG
SPKNO
LFLNG
LFWD
LFRATIO
LFCOL
LFPUB
HABIT
AWNS
HEAD
BLOOM
MATURE
SITE=CF
year=2008
45.4
32.1
17.8
18.3
17.1
44.4
34.1
17.3
16.5
21.4
24.6
33.6
31.9
13.3
40.6
38.8
21.9
17.8
SITE=LP
year=2007
46.9
43.4
8.2
31.5
25.6
35.1
20.8
24.1
25.6
13.1
38.4
7.5
23.3
25.5
26.5
14.2
1.1
0.0
SITE=LP
year=2008
20.8
22.8
12.3
23.3
21.6
33.3
18.1
23.0
18.6
56.4
36.5
37.2
25.6
27.7
54.7
31.2
44.0
13.0
SITE=PU
year=2007
36.5
33.0
28.5
36.5
34.5
18.5
8.8
22.9
14.0
32.5
37.4
29.9
11.9
28.1
54.9
34.7
47.1
28.9
SITE=PU
year=2008
31.2
28.0
13.3
20.9
18.6
32.0
28.3
35.9
16.3
0.2
15.0
3.1
38.6
6.9
45.3
19.8
27.6
9.3
27.4
30.3
12.1
24.9
18.8
24.1
28.4
25.5
18.6
9.3
37.4
4.8
25.8
22.4
42.5
11.9
40.7
4.5
Bluebunch: dry weight
Primarily affected by
precipitation and extreme
minimum temperature
EMT 0.37
MSP 0.32
MAP 0.31
Regression:
1138.87 - 13.51*MWMT + 7.24*TD +
0.05*MAP - 0.16*SHM -3.2*bFFP - 1.91*FFP 0.2*PAS + 7.9*EMT
Bluebunch: heading date
Primarily affected by
beginning of frost free days
and precipitation as snow
bFFP 0.47
PAS 0.21
Regression:
147.07 - 5.34*MAT + 2.6*MCMT + 1.48*TD +
0.53*EMT + 0.03*Eref
Bluebunch: leaf width:length ratio
Primarily affected by heat
moisture index and mositure
deficit
AHM
CMD
SHM
MWMT
MAT
Regression:
0.63
0.59
0.58
0.55
0.53
20.33 + 1.11*MAT + 0.2*AHM
Bluebunch: PC 1
Primarily associated with
plant size
Dry weight 0.94
Crown width 0.93
Regression:
17.12 + 0.02*TD - 0.02*SHM + 0.47*EMT
Bluebunch: PC 2
Primarily associated with
phenology
Bloom
0.87
Heading date 0.82
Regression:
3.37 + 0.02*TD - 0.007*SHM - 0.02*FFP
Bluebunch: PC 3
Primarily associated with leaf
ratio
Leaf ratio
Regression:
0.73
-2.07 - 0.004*PAS + 0.004*CMD
Bluebunch: Comparisons of native collections with
cultivars
Noncultivars
Anatone
Goldar
P-7
Whitmar
Secar
Dry wt
76
151
154
160
131
201
Crown width
8.4
9.2
9.0
9.8
9.4
9.7
Inflorescence no.
88
199
172
159
168
201
Leaf form
38
39
32
41
34
42
Leaf color
3.1
3.3
3.5
3.6
3.7
3.4
Leaf pubescence
4.2
4.3
4.2
4.0
4.2
2.9
Plant form
6.3
5.7
5.5
5.7
5.4
6.5
Awn length
4.6
1.9
3.9
1.7
1.1
6.2
Heading date
137
136
1.40
139
138
142
Bloom date
152
152
150
154
157
154
Maturity date
191
190
192
192
189
192
Trait
Cultivars are more vigorous in test environments. Selection works.
Cultivars are variables w.r.t. leaf width, phenology, and awn length.
Secar is most different = Elymus wawawaiensis
Bluebunch: climate change
Current climate
2050
Bluebunch: recommended seed zones
early
early
late
late
small
Intermed.
large
1
red
brown
13.4%
3a
yellow
24.7%
6
light blue
4.0%
wide
none
3b
7
light
dark blue
orange
2.6%
1.2%
narrow
2a
dark
orange
4.4%
4
light
green
17.6%
8a
light
purple
1.7%
wide
2b
orange
2.5%
5
dark
green
17.1%
8b
dark
purple
11.0%
narrow
Conclusions
• High levels of population variation in many traits
• Correlations with climate are fairly strong
• Relationships make sense from an adaptation perspective
– Larger, more vigorous plants come from populations located in wetter, mountainous
regions without large seasonal temperature differentials
– Plants with later heading and anthesis dates come from populations located in
colder, less arid climates
– Plants with narrow leaves come from populations located in warmer, more arid
climates, particularly in the summer
– Plants with more leaf pubescence come from populations located in drier climates
• Thus, strong evidence for adaptively significant genetic variation
• Seed zones are proposed that follow these gradients in
traits/climates, but be cautious about transfers between Level 3
ecoregions.
Climate matters
Prairie Junegrass (Koeleria macrantha)
Field Sites: NRCS Plant
Materials Center, Corvallis;
Powell Butte
127 populations
www.illinoiswildflowers.info
Katherine Bolin: www.wildflower.org
Wikicommons: www.wikipedia.org
Prairie Junegrass (Koeleria macrantha): ANOVA
F-value
Trait
Year
Site
pop
Year*Site
Site*pop
Year*pop
head
7058.15***
3610.78***
6.45***
13722.3***
3.63***
3.4***
bloom
282.18***
3351.85***
4.76***
1463.21***
1.87***
1.36**
mature
163.4***
2209.93***
1.18
404.87***
1.18
1.06
inflornum
1068.1***
19.44***
4.11***
2074.04***
2.63***
2.13***
culm_cm
586.3***
122.75***
2.19***
421.62***
2.57***
1.13
inflor_len_cm
500.83***
9.64**
3.69***
410.91***
2.51***
1.38**
lfratio
2780.53***
3.98
4.62***
300.85***
1.48*
1.29*
max_wcm
245.86***
53.71***
1.63**
13.8***
1.19
1.03
form
258.83***
428.15***
2.46***
526.64***
2.19***
2.41***
biomass_g
2383.21***
70.69***
9.97***
368.53***
9.15***
6.68***
* < 0.05
** < 0.01
*** < 0.001
Prairie Junegrass (Koeleria macrantha): Percent
trait variation due to population of origin
CRV
PB
Trait
2009
2010
2009
2010
head
18.7%
32.0%
31.9%
32.8%
bloom
1.4%
25.5%
35.4%
39.2%
mature
0.9%
29.4%
2.3%
2.4%
inflornum
22.2%
17.0%
7.2%
13.0%
culm_cm
3.4%
22.6%
3.8%
7.8%
inflor_len_cm
23.5%
11.3%
7.6%
13.3%
lfratio
11.1%
5.3%
14.4%
10.1%
max_wcm
26.9%
0.7%
7.4%
8.1%
form
16.9%
15.9%
11.2%
7.3%
biomass_g
38.0%
52.0%
15.0%
10.9%
Prairie Junegrass (Koeleria macrantha): Percent
trait variation due to population of origin
Trait
MAT
TD
MAP
Biomass_g_crv09
0.22
-0.40
0.28
Biomass_g_crv10
0.16
-0.32
0.33
Biomass_g_pb09
0.18
-0.43
0.11
Biomass_g_pb10
0.05
-0.11
-0.31
Head_crv09
0.27
-0.17
0.19
Head_crv10
0.22
-0.13
0.20
Head_pb09
0.22
-0.25
0.35
Head_pb10
0.27
-0.44
0.46
lfratio_crv09
0.37
0.20
-0.18
lfratio_crv10
0.24
0.33
-0.22
lfratio_pb09
0.40
0.26
-0.20
lfratio_pb10
0.35
0.29
-0.37
Bottlebrush squirreltail (Elymus elymoides)
114 populations, 7 variety/pre-variety, 3 field sites (Central Ferry, Powell Butte, Reno)
Collaborators: Matt Horning, RC Johnson, Elizabeth Leger
Thanks to many folks who
contribute to seed collections