. l. ...
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lPACTIlFTI<C
§OUTHWJE§T
FORJE§T & RANGJE JEXJPJERJIMJENT §TATJION
19 6 7
P.O . Box 245
Berkeley.
, Winter Injury '
California
94701
ABSTRACT:
White fir seeds collected
from 43 sources were sown at the
Institute of Forest Genetics.Pla~er­
ville. Calif .• in 1963. ObservatIons
made 3 years later showed that s~ed­
lings from northern sourc~s sustaIned
more winter injury than dId southernorigin seedlings. Seedlings from low
elevations were less severely dam~ged
than seedlings from higher. elevations
in the same general localIty.
Among White Fir Seedlings
unusual pattern in seed source study
M.THOMPSON CONKLE
W.
J .L.HAMRICK.
J. LIBBY
III
RETRIEVAL TERMS:
Several weather conditions are
known to produce winter injury. Foliage, buds, and cambium can be injured
by early fall frosts, by unseasonally
cold winter temperatures, by re-radiation of sun l ight within snow or from
a snow surface, by winter desiccation,
and by late spring frosts.
Reports of winter injury in plantations or nurseries have generally lead
to the expectation that--within species--plants from lower latitudes or
lower elevations wi l l be more susceptible than those from higher latitudes
.
1
or higher elevatlons.
In 1962 a cooperative study was
begun between the U.S. Forest Service,
University of California at Berkeley,
and Michigan State University to examine the within-species variation of
white fir (Abies concolor [Gord. &
Glend.] Lind!.). In February 1966, we
observed damage in the form of needle
kill on many of the seedlings (then
2-1 transplants). The source-tosource variation in injury was found
to be greater than the seedling-toseedling variation within a source.
Furthermore, the pattern of injury
with regard to latitude and elevation
of seed source was unexpected.
1Clauson
Keck
and Hiesey 1941; Eiche 1966;
Ernstso~ and Badders 1948; Sto~ckeler 1948;
Stoeckeler and Rudolf 1949; WrIght 1962.
F orest
Serv i ce
-
U.
S.
Abies concolor; leaf
damage; low temperature; .s~ed sour~es;
seedling damage; winter InjUry. LIne
Project: 1401.
Materia l s And Methods
Seeds were collected from 43 sites
from throughout much of the white fir
range (fig. 1). One seed source col lection was made from a single seed
tree, a few collections consisted of
seed from two to six trees, while most
collections came from seven or more
parent trees within a local area. The
seeds were sown in the Institute of
Forest Genetics nursery, Placerville,
Calif., in spring of 1963. In spring
of 1965, two IS-tree rows of each collection were transplanted into each of
four replications within five nursery
beds (table 1).
This systematic positioning within
replications precluded the unbiased
use of statistical analysis. The repetition of the injury pattern over the
four replications in transplant beds,
and the predictability of damage in
other beds (which did not contain all
43 sources), however, gave us considerable confidence that differences in
injury between seedlings of different
sources were real.
The injury produced a general browning of needles that was followed by
needle drop within 4 weeks. No seed lings were killed. Needle injury was
scored on a four - point scale:
Department
of
Agriculture
.
..
.
.i·
,.
----- --'-
BF
o
'- '- '-
oAR
oAS
BB
oAT
ol
50
200 Miles
c ==~I====±====±==~I
o
... ......
............
o
Figure 1.--Seed collection sites and average injury value for white f i r s e edlings a t
the Institute of Forest Gen e t i cs, Pla ce rvill e , Ca lif .
Alphabe ti c s y mbol s a r e IFG
s eed lot c odes.
•
o
None
slight (up to but not exceeding one-third of the total
number of needles were injured)
2 . .. moderate (one-third to twothirds of the needles were
injured)
3 ... severe (more than two-thirds
of the needles were injured),
injury value for the lower collection
(AK) was the smallest of the four.
Collection AL, although from an elevation intermediate between AK and AN-had the highest injury value. However,
the percentages of seedlings with no
injury for these four sources decreased
as elevation of co llection increased .
1
Twenty seedlings of each collection
in each replication were scored. An
average injury value based on 80 seedlings was calculated for each collection .
Resu lts
Contrary to our expectation, the
seedlings bf northern origin were the
most severely damaged ~able 1). Ihose
most severely damaged were from lot R
collected at 5,000 feet in the Ochoco
Mountains of central Oregon. In Oregon and California, from north to
south, the injury values decreased in
a clinal pattern. Few seedlings from
sources south and east of the Sierra
Nevada were injured, and in three cases
no detectable injury was noted on any
of the 80 plants scored.
Each of the 43 population samples
contained a few seed l ings with no detectable winter injury (table 1). Collection R, for instance, with 57 percent severely -injur ed seedlings,
included 7 of 80 seedlings apparently
uninjured.
Pairs of collections--one from a
higher e levation than the other-were made within five relatively limi ted geographical areas. In all five
cases, seedlings from the lower elevation had less injury than seedlings
from the higher e l evation (table 1).
Collections AK, AL, Mvl, and AN
transect El Dorado County in California at about the same latitud e but
different elevations. The average
Injury was negati ve ly correlated
with 2-year height 2 of tr ees from Oregon, northern California, and western
Nevada. Seedlings from the Oregon collections were the smallest and most
severely damaged. This correlation,however,
does not hold for the eastern collections and the southern California collections , which varied greatly in 2year height, but were nearly free of
injury.
Discussion
Winter injury to plants has been
attributed to three main causes: (a)
intrace llular freezing (very uncommon
in nature); (b) freezing and thawing
of extracellular water; and (c) dehydration sometimes referred to as
physiological drought (Scarth 1944) .
These injuries can be related to the
physiological condition of the plant.
A winter -hardy plant can survive temperatures below the freezing point of
water provided water is held by the
proteins of the cellular protoplasm
(Levitt 1956). On the other hand,
plants lacking the abi lity to ret ai n
"bound" water are subject to freezing
of extracellular water and to plant
dehydration (Peace 1962 ; Weav er and
Clements 1929).
Nursery records of maximum and
minimum ai r temperatures, soil temperatures at depths of 1 anq 4 inches,
precipitation, and wind were examined
for the 5-month period preceding observations of injury symptoms. During
2Hum :- ick . .f . L . Geog r aph i c va riat'~;n in white
fir . 1966 .
(Unpublished master ' s thesis on
file at School of Forestry , University of
California, Be d celey ,)
-3-
Table 1. ~ -Wint er injury amon g whi te fi r seed sources , by co ll ection si te
IFG
seed
lot
Seedlings showing
Latitude
Longitud e
Elevation
No
injury
I
Sli ght
injury
Feet
0
N
45 17 ' N
~J 11
44 "48'
118°31' W
118°01'
118°11'
120°45'
1
120°45
c
AI
44°47 1
1
44°30
44°30 1
44°18 ;
43°00'
43°00'
42°47'
42°01 '
42 °05'
42°21 /
42°29 !
41 °28 '
41 °30 1
40°45 '
40°42'
40°27;
40°25'
39°29'
AJ
39°54 '
121:°05;
NT
38°33 '
38°54 '
1
38°51
38°48'
37°44'
36°56'
34°50'
120°26:
120°23 1
120°00'
120°14'
118°55'
119°02'
119°10'
34°21 '
34°10'
1
33°47
33°20'
32 ° 57 '
1
36°15
39°02 '
39°02'
34 °26 '
117°40'
117°05'
116°45 '
116°52'
1
116°35
115°35 '
114°15 '
114 °58 1
112°25 '
BF
34 °06 '
35°00 1
39°20'
39°15'
39°16'
a-:;
40 2S
110°56'
111 °30 '
120°52'
120°57'
111 ° 27 '
111 ° 44 '
~l~j
S
~J -ll
V
W
X
Y
Z
Ai>.
AB
AC
AE
:J-Y
AI.,
AM
AN
AO
AP
AQ
AR
AS
AT
AU
AV
AW
AX
AY
l3A
BE
OC
:J-Y
0
0
122 05
i
121 ° 50 '
121 ° 50 '
122°04'
123°29 '
122°47 1
121°15 i
.
f
,.'
120°55'
121 ° 29 '
120°15'
0 i
123 40
121 °46 .'
121 ° 05 '
121°06 '
122°441
4 , 400
4 , 600
5,000
4 , 000
5 , 000
4 , 000
4 , 500
, 5 , 500 ..
4 , 500
5 , 200
5 , 600
5 , 000
7 , 000
5 . 400
- 6 , 000
5 , 400
4 ,\800
5 , 800
6 , 500
6 , 1006 , 200
3 , 7005 , 000
4 , 200
5 , 500
6 . 800
6 ,900
7 , 600
7 , 200
6 , 1008 , 500
6 , 200
5 , 700
5 , 500
5 , 100
5 , 600
8 , 400
7 , 500
9 , 000
6 , 9007 , 000
7 , 500
7 , 500
7 , 700
6,600
7,800
5 , 600
.
I
..
Moderate
injury
I
Severe
injury
Average
inj u r y
valu e ll
P e rc en t
10
14
12
19
9
10
39
30
20
35
45
56
42
78
56
50
91
80
55
46
21
16
13
11
18
23
25
12
11
16
30
30
15
24
21
8
10
24
16
19
29
31
34
23
36
22
26
35
35
32
11
18
6
14
24
1
9
20
33
50
41
44
36
57
36
16
19
33
19
7
3
10
1
6
5
0
1
1
5
2 . 09
1. 98
2 . 06
1. 88
2 . 29
1.99
1.16
1.34
1080
1. 38
74
8
17
1
.46
89
80
79
74
81
79
99
6
4
5
20
6
15
1
3
10
15
5
13
4
0
2
6
1
0
. 19
. 43
.39
, 34
. 31
.30
. 01
94
94
93
99
99
98
100
100
100
5
2
7
1
2
0
0
0
1
4
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
. 08
10
, 08
01
. 01
.02
. 00
. 00
. 00
99
84
43
47
99
99
0
14
15
24
1
0
1
2
26
19
0
1
0
0
16
10
0
11
0
1
0
:2
0
LOO
60
. 95
31
. 70
. 84
, 10
. 31
. 68
.96
..;
;',-
.01
.19
1.16
. 91
. 01
. 01
~~"
lSca l e used , 0"'00 injury ; l=sllght injury ; 2=moderate injury ; 3",severe in j ur y
2Brackets enc l ose col l ec t I ons t aken at a bout th e same locat i ons but at different elevations
3Brackets enc los e tr ansect taken along U. S. 50 a t low, medium , and high elevations ; on th e
wes t side of the Sie~ra , and from a n east- si de population,
-4-
the 10 days before visible injury appeared, the maximum daily air temper··
atures ranged from SoC. to 14°C . and
minimum temperatures from _2°C. to
2°C. Records of soil temperatures
indicate the soil was not frozen during this period. These air temper atures are not in the range nor of the
length that Daubenmire (1957) reported
for winter damage to native and introduced trees and shrubs in Wash i ngton
and northern Idaho.
The period preceding the appearance of visible injury was also characterized by high winds and no
precipitation. If the minimum temperatures did not cause the damage, dehydration by high winds seems a second
plausible explanation of the cause of
injury.
The pattern of injury is similar
to that observed for jack pine (Pinus
banksiana Lamb.) and--to a les ser extent--for red pine (P. resinosa Ait.)
planted in Texas. 3 All of the jack
pine froze at temperatures only
slightly below freezing, which occurred after warm weather in late winter
had apparently triggered the pines
into growing. Wright (1962, Ch. 6)
points out that northern provenances
of widespread species are often more
susceptible to late spring or early
autumn frost , although they are more
resis tant to damage by winter cold.
White fir has two geographic races
that have been given taxonomic recognition (Lamb 1914; Sudworth 1916).
One race is found in the Rocky Mountains and westward into Arizona and
southern California. The second occupies the Sierra Nevada and extends
through northern California into
southern Oregon.
In the Cascade region of northern
California and Oregon, another taxonomic problem occurs. A. concolor is
in close proximity with a closely related species--grand fir (A. grandis
[Dougl.] Lindl.). Many authors (Lamb
1914 ; MUller 1939; Maul 1959) have
commented on the difficulty of telling the two species apart in this
region.
Artificial hybridization
is possible between these two species
(Gathy 1957; Scheplitz 1956).
Seedlings in the collections from
the Rocky Mountains, Arizona, and
southern California showed no appreciable damage.
Seedlings from
the Sierra Nevada, northern California, and southern Oregon sources had
slight to moderate damage, with
greater damage in the more northern
sources. The pattern of winter injury conforms to the pattern of
growth differences that have been
observed. 4 The collections from the
east-central Oregon and northwestern
California populations which tend to
be int ermed iate between A. concolor
and A. grandis showed moderate to
severe damage. No A. grandis sources
were included, and the response of
that species is unknown.
Weather records from the Institute
of Forest Genetics revealed no unusually severe conditions in fall of
1965. Furthermore, the seedlings
appeared healthy in January, had
brown needles in February,
and had shed the injured needles by
early March. This seems to indicate
that injury occurred in mid-winter
rather than in fall.
We found differences between seedlings from paired collections at
about the same location but from different elevations. Though some coll ec tion pairs differ ed only 400 to
700 f eet in elevation, the injury was
consistently greater for seedlings
from the higher elevation collections.
3personal correspondence with B . J . Zobe l ,
professor of forestry , North Carolina
State University , Raleigh , N. C. , March
1966 .
4Eamrick ,
-5-
J . LOp . ci to
Ernstson, M., and Hadders, G.
1948. Skadegorelse a granplantor
genom utterkning unter varvintern
194 7.
(Damage to Norway spruce
plants by drying out in the winterspring period of 1947.) Svenska
SkogsvaardsForeningens Tidskr. 46
(5):310-322 .
This t rend is consistent with the
greater injury to se edlings from hi gher l atitudes.
The Institute of Forest Genetics,
at an e levation of 2,700 feet, is near
th e lower limits of white fir in that
area . A 1965 out-planting of seedlings from thi s s tudy at Camino, only
6 miles away, anu 500 f eet hi gher in
elevation, did not show injury of the
type observed i n the nursery. Other
1965 plant ations of similar seedlings
at four other sites throughout northern Ca liforni a, and within the nativ e
range of white fir , did not show this
inj ury .
Gathy , P.
1957. A propos de l 'hybride naturel
Abies concolor (Cord .) Engelm. x
Abies grandis Link. Silvae Genetica 6(6):186-190.
Lamb, W. H.
1914. A conspectus of North American
firs (exclusive of Mexico) . Soc .
Amer. Foresters Proc. 9:528 -5 38,
illus.
Whit e fir is a favored species for
Christma s tree plantations at sites
outside the species' native range.
Injury of the type noticed in the nursery may discourage the us e of seed
from Oregon and northern California
in such plantings.
Levitt, J .
1956. The hardiness of plants. Amer.
Soc. Agron. 278 pp.
Maul, D. C.
1958. Silvical characteristics of
white fir. u.s . Forest Servo Calif.
Forest & Range Exp. Sta. Tech. Paper
25, 22 pp ., illus.
Th e physiological basis of the injury and the regularity with which it
will occur in the future is unknown .
But the injury symptoms show an underlying variability in the physiology or
anatomy of the seedlings. This variability is strongly related to the geographic origin of the seed.
MUller, K. M.
1939. Abies grandis and its climatic
races . Mitt. dt. dendrol . Ges.
136 pp.
Peace, T. R.
1962. Pathology of trees and shrubs
with special reference to Britain.
753 pp. London : Oxford Univ. Press.
Lit era ture Cited
Clausen , Jens, Keck,D.D .,and Hiesey ,W.M.
1941. Regional differentiation in
pZant species. Amer. Natur . 75:
231-250.
Daubenmire, R.
195 7. Injury to plants from rapidly
dropping temperature in Washington
and Northern I daho. J . Forestry
55( 8):581-5 85.
Scarth, G. H.
1944. Cell physiological studies of
frost resistance: a review. New
Phytol. 43:1-12.
Scheplitz, B.
1956. Uber einen naturlichen AbiesBastard. Morphologische und holztechnologische UnterBuchung an A:l:'tbastarden von Abies c.onc6loY'x Abies
grandi,f3 . .0. nat ural Abies hybrid. Investigation into morpho lo gy and wo od
technology of a species hyb rid of
A. concolor x A. grandis. ) Z. Forst genet . 5(3) :7 1-79. (Forest. Abstr.
vol. 17, No. 3646.)
Eiche, Vilhe lms.
196 6. Cold damage and plant mortal ity in experimental provenance
plantations with Scots pine in
northern Sweden. Studia Foresta1ia Suecica 36:1-218, illus . Stockholm : Skogshogskolan.
- 6-
trees of the Rocky Mountain region.
Stoeckeler, J. H.
1948. Recovery of winter-injured
conifers. Arner. Nurseryman 88:9.
U.S. Dep. Agr. Bull. 327, 43 pp.,
illus.
Stoeckeler, J. H., and Rudolf, P. O.
1949. winter injury and recovery of
conifers in the upper midwest. u.s.
Forest Servo Lake States Forest
Exp. Sta., Sta. Paper 18, 20 pp.
Weaver, J. E., and Clements, F. E.
1929. Plant ecology. 520 pp. New
York: McGraw-Hill Book Co., Inc.
Wright, J. W.
1962. Genetics of forest tree improvement. F.A.O . , Forestry and For-
Sudworth, G. B.
1916.
The spruce and balsam fir
est Prod. Study 16, Rome, 399 pp.
The Au tilors _ _ _ _ _ _ _ _ _ _ _ _ _ _ __
M. THOMPSON CONKLE is a geneticist doing
research on the genetics of western conifers,
wit h headquarters in Berkeley and Plac ervil le ,
Calif. A 1962 genetics graduate of No rth
Carolina Stat e University , h e joined th e Station's r esearch staff in 1965 . W. J. LIBBY
is associate professo ~ 0f forestry at th e
University of California " Be rkeley . He holds
an MS. degree (1959) in tree physiology and a
Ph.D . degree (i961) in genetics-··both from the
University of California . J . L . HA~ffiICK is a
g raduat e student in the De partmen. of Gen e tics ,
Universit y of Califo r nia , Be rkel ey . He earned
an MS. degree in forestry (1966) from th e Uni versity of California .
-7-