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Geographic
Fariationm
PonderosaPine Leader Growth
BY
JAMES
W.
HANOVER
Abstract.Growthof the shootaplcesof 91 treesin a 4S-year-old
Pinusponderosa
Laws.provenance
testwasmeasured
periodlcally
with a transit.Analysis
of the
measurements
led to the followingconclusions:
(1) 19 racesof ponderosa
pineplanted
nearPriestRiver, Idaho,showed
phenologica],
morphological,
or physiological
variationin sixcharacters:
dateo.fbeginning
growthin the spring,dateof endinggrowth,
totalseasonal
elongation,
duratlo3q
of growth,lengthof dormantterminalbud,and
rapidityof growth;(2) thevariation
described
is continuous,
although
it wasnot
possible
to asse.ciate
an environmental
gradient
to thevariation
pattern;(3) all trees
except
thelocalKaniksu
source
achieved
theirmaximum
surgeof growthduringthe
same7-dayperiod,May 20-27,andthisresponse
closely
paralleled
temperature
fluctuation;
(4) sim?le
correlations
among
the19progenies
between
totalseasonal
elongation and eitherbeginning
date,relativerapidity,or durationof growthwerenot
significant.
Correlation
between
dormant
terminal
budlengthandtotalseasonal
elongationwasstrong.Also.,
budlengthwaspositively
correlated
withrelative
rapiditywhich
wasnegatively
correlated
with duration
of growth.
FEW
CRITICAL
OBSERVATIONS have been
made of intraspecific
variationin the seasonalactivityof shootgrowthon tall trees.
A 45-year-oldprovenance
testof ponderosap•ne(Pinus•3onderosa
Laws.) provided
a good sourceof material for measuring
growth of mature tree apiceswith some
environmentalcontrol The objectivexvas
to determinewhetherprovenances
of ponderosapineshowvariationin date of beginningor cessation
of leadergrowth,
rationof growth,rapidityof growth,total
seasonal
elongation,
andlengthof dormant
(1936) and Morris et al. (1957) observedtimesof lateralbudbursting
on 18and 40-year-old Douglas-fir provenance
plantations.They suggestthat this characteristicis under stronggeneticcontrol.
I•ens-Moller (1957) attemptedto determinethe degreeof geneticcontrolover
time of bud burstingby measuringit on
about 700 Douglas-firseedlings,repre-
sentingsevenelevations
from 60 to 4,000
feet. The measurements were made after
the seedlings
had overwinteredin Corvallis,Oregon. His resultsindicatedsig-
terminal bud. Measurements indicate that
nificant differences betxveen certain sources
each of thesegrowth characteristics
vary
significantly
withinthe species.
The variation is closelyassociated
with seedsource
in dateof budburstingthe followingspring
location.
Pest Studies of Shoot Growth
Bud burstingof Douglas-fir,Pseudotsuga
menzlesli(Mirb.) Franco, has received
considerable
attention.Mungerand Morris
86 / ForestScience
and "some correlation" b.etween this char-
acteristicand elevationof seedlingsource.
Chingand Bever (1960) observed
dates
of bud bursting of Douglas-fir nursery
The authoris a geneticist,Intermonntain
Forestand RangeExpt. Sta., ForestService,
U.S. Dept.Agric.,andisstationed
at Moscow,
Idaho.
seedlings
representing
14 sourcesvarying
in latitude(42ø20' to 50ø30') andelevation(100 to 4,100 feet). They found
close relation between latitude or altitude
of sourceand date .of bud bursting,althoughthe southernmost
sourcewas significantlyearlierthan other p.rovenances.
Vaartaja (1954, 1959) hasshownpatterns of geographicvariationin growth
responsefor many tree species.Kriebel
(1957) obtainedevidenceof clinal geographic variationin dates when sugar
maple(ricer saccharum
Marsh.) from 18
seedsources
commenced
growth. IrgensMoller (1958) demonstrated
geographic
variationin time of height growth cessation amongDouglas-firseedlings.Critch-
field (1957), workingwith lodgepole
pine
(Pinus contortaDougl.) seedlings,
associated a continuousvariation in growth
durationwith changes
in altitudeand latitudeof seedsource.Langlet (1959) analyzed Wright and Baldwin's (1957)
data on the growthrate of 46 Scotchpine
(Pinussylvestris
L.) provenances.
He interpretedthegeographic
variability
in height
incrementas continuous
and relatedto day
lengthandtemperature.
Baldwin(1956)
measuredterminal growth during two
seasons
on 9- and 10-year-oldtreesof a
Europeanlatch (Larix declduaMill.)
provenance
plantation
in New Hampshire
but found no correlation between either
•ltitude or latitude of seed source and
all speciesoccurredabout the sametime.
Tryon and Finn (1937) notedthiswhen
theyobservedgrowthcurvesfor red pine
( Pinusrexinosa
Ait.), whitespruce( Picea
glceuca(Moench) Voss), and European
larchin New York State.Friesner(1942)
notedthesimilarityin datesof peakgrowth
within treesof Pinusbankdanaand P. sylvestrisL. Fowells(1941) comparedseasonalheightgrowthof sixconifersgrowing
within a 10-acre area in California and re-
porteddifferencesin beginning,duration,
and rapidityof growth.
This brief reviewof pastwork indicates
clearly that the genetic makeup.of the
source material
and a number of climatic
factorsmay interactto producegeographic
variationin shootgrowth characteristics.
Through naturalselection,
populations
becomeadaptedto each local environment.
Sometimes
the adaptation
canbestatistically
correlated with environmental
factors. In
othercasesthisis not poss•lebecause
interpopulationmigration, limited populati.on
sizes,and slow geneticresponse
to selection pressuretend to obscurephenotype
correlations.Under growthchamberconditionsor by appropriatesamplingtechniquesand experimentaldesign,the influence exerted on growth by any one
environmental
factormightbedetermined;
but this would be difficult for the species
and growth characteristics
being considered here.
•m.ountof currentheightgrowth or mid-
pointsof growthcurves.He observed
that:
(1) all sources
completed50 percentof
their growth at approximatelythe same
date,(2) beginning
dateof growthvaried
as greatlywithin as betweenprovenances,
and (3) air temperaturewas associated
with bud bursting. Kienh.olz (1941),
Brown (1915), Friesnet (1942), McMillan ( 1957), andothershaveattributed
the inceptionof bud growth largely to
temperature effects within certain photo-
periodic
andgenetic
limitations.
Severalauthorshave comparedterminal
growth betweenspeciesgrowing at one
2tocality.The peakof seasonal
growth for
Experimental Materials
Seedor seedlings
representing
22 provenancesof ponderosa
pine were plantedat
the Priest River ExperimentalForest in
northern
Idaho
from
1911
to
1917.
Weidman (1939) hasfully described
the
climatenear the pointof originof eachof
the 22 seedsources
and hasalsogenerally
described
the geographic
areasof the seed
sources(see also Table 1 and Fig. 1).
The seedsources,
or progenies,
fairly well
representthe broad range of ponderosa
pine in the westernUnited States.Since
detailsof plot establishment
were well devolume9, hum,herl, 1963 / 87
T.4BLE 1. Geographic
locationof ponderosa
pine seedsources
represented
at Priest
River Experimental
Forest.
•
General geographicarea
and national forest
Elevation
Latitude
Longitude
Feet
North Pacific
Siskiyou
2,000
42 ø
05'
123ø
40'
Boise
5,500
43 ø
30'
115ø
00•
Payette
5,000
44-0
30'
116
North
Plateau
Whitman
5,000
44 ø
38'
118ø
Umatilla
3,500
46ø
00'
117ø
25'
30•
Bitterroot
Bitterroot
Bitterroot
4.,000
5,000
7,200
46 ø
46 ø
46 ø
00'
00'
00'
114ø
114ø
114ø
20'
20'
20'
Lolo
Kanlksu
Colville
3,000
2,600
2,700
47ø
48 ø
48 ø
10'
20'
40'
114ø
116ø
119ø
•0'
50'
00'
7,100
35ø
10'
Ill ø
50'
8,000
35ø
40'
105ø
30'
Roosevelt
Ashley (Central Plateau)
Custer
8,000
7,500
3,200
40ø
40ø
45ø
30'
40'
30'
105ø
109ø
104ø
40'
40'
00'•
50'
South Plateau
Coconlno
Santa Fe
East of Continental Divide
Helena
4,500
46 ø
30'
111o
Harney (Black Hills)
5,000
43ø
40'
103ø
30'
San Isabel
8,000
38ø
00'
105ø
00•'
•Elevation
2•380feetl latitude48ø20'• longitude116050
'.
scribedby Weldman they are not elaborated here. Originally each progenyplot
was composed
of either 100 or 50 trees,
but mortalityof varyingdegreeshas reduced the number of trees remaining to
10 to 50 per plot. All treesfrom one of
the originalsources(Mt. Shasta,California) were eliminatedentirelyby a severe
temperaturedrop in December 1924
(Kempff 1928). Two sources
of questionableoriginalsowere omittedfrom this
study.
The Priest River test of seed sources has
receivedconsiderable
studysinceits establishment. Individual progenyhave been
previously
examined
and compared
for differences in vigor and frost resistance
(Kempff 1928); external and internal
foliagecharacteristics
(Weldman 1939);
total heightand diametergrowth (Weldman 1939, Squillaceand Silen 1962);
and seasonof cambialgrowth (Daubenmire 1950).
88 / ForestScience
Methods
and Definitions
Methods of measure'men:t.
Seasonalactivity of shootapiceshas never been describedfor these progenies. In March
1958 five dominant or codomlnant trees
rangingfrom 25 to 65 ft in heightwere
selected for measurement from each of the
19 plots.Their initialterminalbudlength
and subsequent
heightgrowthwere measuredthroughout
the course
of thegrowing
season.
The convenient method of direct meas-
urementabovea constantpoint, suchas a
pininserted
at thebase.oftheterminalbud•
was impractical
because
of the tree height
and top flexibility. Therefore, the indirect
method of angular measurementswith a
transitwasadopted.Nine fixedinstrument
stations were established from which the
tipsand basesof all selected
treescouldbe
sightedclearly. The distancefrom each
tree to a station was measured. Graduated
sticks were nailed to the base of each tree
to permitadjustmentfor heightof instru--
ment at each remeasurement.
The onlydifiqculty
encountered
in using
this method was causedby wind. Any
shghtbreezeconsiderably
reducedthe accuracyof an observation
l consequently,
all
measurements
weremadeonlyduringcalm
periodsof the day, usuallyearly morning
or evening. Error in measurement
result•ng from the exper/mentaltechniquedid
not exceed --+0.03 ft, well within the
precisionnecessary
for statisticalvalidity.
This value was determinedby averaging
I
FmURE1. Climaticregions
of ponderosa
pinerange(northof Mexico):(I) NorthPacific;
(2) northplateau;(3) centralplateau;(4) southplateau;(5) eastof Coutin,ental
Divide; (6) SouthPacific.The brokenline shows
therangeof ponderosa
pineaccording
to Sudworth
(1917) andMunns(1938). Localities
fromwhichseedusedin experiment
wasderivedareindicated
by blackdots.The experimental
siteis located
at the Kaniksu
dot. (From Weldman19.39)
volume9, numberI, 1963 / 89
the variationbetweenreadingsobtained to theseheightswere usedin the analyses.
from successive
measurements
duringthe
Mirov et al. (1952) useda method
based on lines of intersection of observed
periodwhennogrowthwasoccurr/ng.It
includes
errordueto instrument
resetting, weekly increment values and average
residual
treeleanaftera wind,andhuman
weeklyincrementover the entiregrowing
error.
Sincethe tipsof four treesdiedduring
the observation
period,completemeasurements were obtainedon only 91 trees.
Treeswere measured
every2 to 4 days
beginning
April 2. Fr.omMay 13, when
all treeswere growing,until July 8,
measurements
were taken about every 7
days.Thereafter,
untilOctober
4, longer
period. Others have suggested
obtaining
the beginningand endingdatesby constructinggrowth curvesfor eachtree and
determining
thedesired
pointson thecurve.
Dependingon the natureof the data,any
of thesemethods
may be appropr/ate.
Duration of growth is the number of
dayselapsingbetweenthe 5 percentand
95 percentdates.Total elonga'tion
is total
intervals
between
measurements
sufficiently increment in feet between the beginning
accountedfor later growth and new bud
and endingbasedates.Bud lengthin feet
formation.
Definitions.The beginning
and ending
datesof growthweredetermined
bycomputingthe pointsof time (expressed
as
consecutive
daysfrom January1) at which
5 percentand 95 percentof the totalseasonalgrowthhad beencompleted.This
was necessary
because
of the difficultyin
pin-pointing
actualgrowthbeginning
and
cessation.
Early growthseems
to besporadicat firstbecause
of dailyweathervariationsand because
smallperiodicincrements
may beobscured
by error in measurement.
Calculation
of thesebasedates(.or 0 percent and 100 percent)is necessary
to
arrive at the 5 percentand 95 percent
points.The beginning
basedate height
was taken as the averageof the first and
all successive
measurements
up to one
showinga difference
of greaterthan 0.05
ft betweentwo consecutive
readingsfollowedby no successive
readings
differing
by lessthan 0.05 ft. Ending basedate
heightwasdetermined
by takingthe average of successive
measurements
not differing by more than 0.10 ft after encountering a difference of less than 0.10 ft
between
two
successive measurements.
Rather than 0.05, 0.10 ft was used to
adjust for the increasedtime betweenobservationsat the closeof growth. Once
the beginningand endingbasedateswere
was computedfrom transitreadingsto the
baseand tip of the budson April 1, when
the first heightmeasurements
were made.
Rapidity of growth, as usedhere, is the
percentof the total season's
growth completed within the 2-week periodof most
rapidgrowth. Rapiditymay be thoughtof
in two ways:(1) absolute,
whichis total
elongation
per unit of time, and (2) relative, which eliminates differences in total
elongation
andcompares
growthrate during the grand periodof seasonal
growth.
The latterismoreappropriate
in this study
becausedifferent geneticpopulations
have
differentabsolute
growthrates(Tables2
and 3). Absoluterapiditymay be more
applicablefor most intraracial or intraspecificstudies,sincelessgeneticdiversity
isusuallyencountered.
The periodof most
rapid growth for all the ponderosa
pine
progenies
represented
in thisstudywas the
2-weekperiodin which38 to.58 percent
of their total growth .occurred.Fowells
(1941) considered
relativerapidityin h•s
studyof seasonal
growthof youngponderosapinein California. However, hismethods of calculation were different.
He used
a constantpercentof growth (50 percent)
rather than a constanttime interval, and
he measured
rapiditybytheminimumnum-
ber of days requiredto completeth•s
amountof growth.
found,the5 percentand95 percent
heights Method of analysis.From the data obwerecalculated.The datescorresponding tained,mean valuesfor eachprovenance
90 / ForestScience
TABLE 2. Ranking
ofponderosa
pinesources
bydateof beginning
andending
ofgrowth
and durationof growthfi
Mean beginningdate
Source
Mean endingdate
Day of year•
Kaniksu
114
Ashley
115
Slsklyou
116
Bitterroot7,200'
117
Umatilla
Source
Mean duration
Day of year2
Bitterroot7,200'
Siskiyou
153
155
117
Lolo
Helena
Boise
156
156
157
Bitterroot4,000'
118'
Bitterroot4,000'
Roosevelt
San Isabel
Colville
Lolo
Helena
119
119
119
120
120
Payette
Harney
120
120
Santa Fe
120
Bitterroot 5,000'
121
Boise.
Custer
121
122
Ashley
Whitman
Coconlno
123
124
Santa Fe
169!
Coconino
175J
Source
Days
Boise
Helena
Lolo
35
36
37
Bitterroot7,200'
37
Custer
37
158
Bitterroot5,000'
38
Kaniksu
Colville
Custer
158
158
159
Colville
39
Siskiyou
40
Whitman
40
Bitterroot 5,000'
159
Bitterroot4,000'
40
San Isabel
Roosevelt
160
161
Payette
Payette
Harney
161
162
Whitman
163
Umatilla
164
167'}
41
42
San
Isab
Harney
Roosevelt
Kanlksu
Santa Fe
Umatilla
Ashley
Coconino
aAnytwomeans
notincluded
withinthesameline appearing
to the rightof eachrankingof sources
aresig-
n•ficantly different.
aDaysof the yearnumbered
consecutively
from Januu'yI, 1998.
TABLE 3. Rankingof ponderosa
pinesources
by totalelongation,
terminalbudlength
(April 1958), andrelativerapidityof growth)
Mean total elongation
Sourc.e
Harney
Mean terminalbud length
Feet
0.93
Roosevelt
.98
Rapidity
Source
Feet
Source
Roosevelt
0.11
Santa Fe
Coconlno
Umatilla
37.9
42.3
43.5
Ashley
.I 2
Percent
Payette
1.06
Coconino
.13
Custer
1.08
Colville
.14
Bitterroot7,200'
1.11
Custer
.14
Payette
Harney
43.7
44.6
San Isabel
1.16
San Isabel
.14
Colville
1.22'
I
Harney
.15
Bitterroot5,000'
Bitterroot4,000'
Ashley
45.3
45.6
46.2
Helena
Boise
S•sklyou
1.23!
1.23/
1.24/
Payette
Bitterroot7,200'
Bitterroot5,000'
.15
.16
.16
Lolo
47.1
B•tterroot
4,000'
1.28/
Bitterroot4,000'
.18
Coconlno
Kaniksu
1.29[
1.33I
Siskiyou
Kaniksu
.20
Ashley
1.12
] Santa
Fe.14
.21
Sisklyou
47.3
San Isabel
Whitman
47.6
48.0
Custer
Roosevelt
Colville
48.6)
48.7/
50.3/
Bitterroot
7,200'
Bitterroot
5,000'
1.37
[ Umatilla
.231
Whitman
1.43[
Helena
Lolo
1.55[
Whitman
Umatilla
1.64•
Lolo
.23
.24
.24
Boise
Kaniksu
Helena
51.4/
51.9/
55.0/
58.1J
•Anytwomeansnot included
•vithinthe samellne appearing
to the rightof eachrankingof sources
are slgmficantly different.
were
calculated
for
the traits observed
(Tables 2 and 3). Either individualtree
data, progenymeans,or both were used
for analysisof varianceof eachcharacteristic (Table 4). Bartlett's (1937) chisquaretest for homogeneity
of variancein
volume9, number1, 1963 / 91
TABLE 4. Analyscs
of variance
of characteristicsof leader growth among 19
sources
of ponderosa
p•ne.
Growth characterlstic
t
F-value
Beginning date
Ending date
2.09*
2.80**
Duration
3.05**
Total elongation
1.97'
Bud length prior to growth
Rapidity (relative)
3.54**
2.39**
XBasedon 5 percentand 95 percentpointsof
seasonalgrowth completion.
*p <0.05• *•p <0.01.
vided further analysisof the variation
amongthe 19 progenies.Any two means
(Tables2 and 3) not includedwithinthe
sameline appearingat the right of each
setof dataare significantly
different. For
instance,in beginningdate of growth the
Siskiyousourceis significantlydifferent
from the Custer but not from the Boise.
The data were alsoanalyzedfor simple
correlations between measured attributes
(table 5).
Results and Discussion
TABLE 5. Coefficients
of simple
correlation (r) between
shootcharacteristics
of
ponderosa
pine sources
at PriestRiver.
Character•stlcs
Progenyor seedsourcebasis(n :
Bud length and total elongation
Duration and total elongation
r
Beginningdate and total elongatlon
t
Bud length and relativerapidity
Duration and relative rapidity
r
19)
0.724***
0.133
0.008
0.458*
--0.599**
Beginning date and total
heightof progeny
t
--0.135
Indlv•dualtree basis(n • 91)
Bud length and total elongation
0.405*•'*
Relativerapidityandtotal elongation --0.126
Bud lengthand relativerapidity
0.257*
XBased
on 5 percent
and 95 percent
of seasonal
growth completion.
*p <0.05• **p <0.01• *** p <0.001.
progeny means showed that differences
amongindividuals
withinprovenances
were
nonslgnificant
at the 5-percent
level. This
wastruefor all characteristics
exceptend•ng dateand durationof growth,which
were nonsignificant
at the 1-percentlevel.
That is, the five individual values from
eachpr.ovenance
wereconsistent
enough
to
justifyuseof a single
statistic
forthesource.
In thetwoexceptions,
theindividuals
varied
too muchto permitassumption
that they
werefrom the samepopulations;
but they
The analyses
of varianceshowsignificant
differences
amongprovenances
for all the
growth characteristics
studied. Multiple
range testsshow that no two adjacently
ranked provenancesdiffer significantly,
when arrayedaccordingto magnitudeof
response,
in any of the traits, i.e., under
the conditionsof this study, the variation
is continuous.
The resultsof simplecorrelationanalyses
betweenattributesprovidesomeindication
as to
the
association between
these re-
sponses
and total seasonalgrowth. At
leastduring the year of this study,neither
beginningdate, relativerapidity,duration
of growth, nor ending date appearedto
have any relationto total growth during
the year. A strong positivecorrelation
existedbetweenbudlengthand total elongation. Bud lengthwas alsosignificantly
correlatedwith relativerapidityof growth.
Thus, total elongatlonseemedto depend
more on inherent growth potential than
on an endogenous
rhythmmechanism
that
gave certainindividualsor progeniesan
advantageover poorerperformingones.
Another interestingpoint revealedby
simplecorrelationanalysiswas a highly
significant negative correlation between
relativerapidityand durationof growth.
An effort was made to relate the con-
tinuous variation patterns to geographic
have some value when aberrant individuals
location of the seed sources. W'eidman
were removed.
(1939) and Squillace
and Silen (1962)
established
relationsbetweenmorphology
or height growth of the ponderosapine
Applying
Kramer's(1956) extension
of
Duncan's(1955) multiplerangetestpro92 / ForestScience
progeniesat Priest River and certain cli-
in the upperportionsof the total elongation scale,and among thesethe sources
The manyinteractingfactorsinvolvedand
from higherelevations
begangrowthlater.
the inadequacy
of the originalplot design Sources
representing
geographic
regionsin
makeit difficultto associate
the responses which September-through-June
precipitastudied here with seed source climatic
tion is relatively low, generally began
factors.
growth later and grew less than seed
Pictorial representation
of severalclisourcesfrom areaswhere precipitation
is
matic factorsand growth responses
(Fig.
high for thisperiod. It is interesting
that
2) provides
insight
intocauses
of thevariathe local Kaniksusourcebegangrowth
tion. This methodhasbeenappliedeffecconsiderably
earlier than the two most
southerlysources,Santa Fe and Coconino.
tivelyin taxonomic
studies
.ofracesor speo
c]eswhen multipleattributesare involved
Four sources--Ashley,
Umatilla,SantaFe,
(Anderson1956). However,evenby this
and Coconino--seem
to have a distinctly
methodonly certain trendsare evidenced
longerperiodof growththanothersources.
and no statistical
significance
is implied.
Suchbroadgeneralizations
aboutprogeny
For instance,the easternsourcestend to
performancein relatl.onto seedsourceare
all that these data allow.
groupinto a sectorof the diagramhaving
a low totalelongation
and a late beginning
The period at which each tree and
date. Only the north plateausources
are
progenyachievedtheir maximumsurgeof
matic factors at the seed source localities.
1.70
I.B0
1.50
1,40
.8O
114
April 24
115
25
116
26
117
27
118
28
119
29
120
50
121
May I
122
2
125
5
124
4
Beginning Date of Grawth (Days numbered consecutively from Jan. l,1958)
FIGURE2. Pictoriallze•d
scatterdiagramshowlnff
variationin threegrowthfactorsand threeclimatic
factorsfor ponderosa
pineseedsources.
volume9, num3er1, I963 / 93
TdBLE 6. Weatherat PriestRiver Experimental
Forestin relationto growthof ponderosapine sources.
Temperatures
Date
Cloud
Maximum
Minimum
øF
øF
Precipitation
cover
x
Inches
Week precedingmaximumgrowth:
May 13
May 14
May 15
May 16
May 17
May 18
May 19
63
74
76
78
79
85
82
30
29
35
36
35
33
43
Tr.
0
0
0
0
0
0
7
2
76.7
34.4
0
2.4
20
21
22
23
24
25
26
85
88
88
85
86
85
90
44
55
46
47
55
51
50
0
0
0
0
0.01
0
0
Mean
86.7
49.7
0.00
91
89
73
76
73
69
72
47
52
46
41
42
41
38
0
Tr.
0
0
0.19
0.20
0
77.6
43.9
0.06
Mean
0
1
7
0
0
Week of maximum growth:
May
May
May
May
May
May
May
Week following maximum growth:
May 27
May 28
May 29
May 30
May 3l
June l
June 2
Mean
0
2
1
10
2
4
5
3,4
1
6
9
5
10
4
6
5.9
•0 z cleari 10 z overcast.
growth was related to local temperature.
With few exceptions
the period of most
rapid growth for all trees was the same
7-dayperiod,May 20-27. Only oneprogeny, the local Kaniksusource,averaged1
response
or for any of the othersconsideredhere. However,Munger and Moriis
ined. Temperaturefluctuationcloselyparalleled this growth response.The week
of maximumgrowth coincidedwith day
and night temperatures
higher than those
of the weeks immediatelyprecedingand
immediatelyfollowingthe week of maxi-
(1936), Friesnet (1942), Tryon and
Finn (1937), and Kienholz
(1941)
showedthat thoughyearlyvariationsoccur
in the absolutevaluesfor manyphenologlcal characteristics,different individuals,
races,or species
usuallymaintainthe same
relativepositions
duringsuccessive
years.
All treesbegangrowingin the spring
beforedangerfrom frost had passedand
ceasedgrowing before the beginningof
fall frosts. Also, growth was at least95
percent completedbefore maximum day
lengthsand the July-Augustdry period
were reached. This growth pattern is
mumgrowth(Table 6). The temperature
characteristicof many coniferous trees
pattern of this year could have obscured
whoseshootelongationdepends
partly on
storedcarbohydrates
but alsoon cell elon-
week
earlier.
In
an effort
to relate
an
environmentalfactor to this response,
weather
records at the Priest
River
Ex-
perimentalForestheadquarters
(lessthan
•-mile from the plantation)were exam-
some differences between sources for this
94 / ForestScience
12:307-310.
gationand divisionin budsformed dur/ng
the previous
year (Sacher1954).
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volume9, number1, 1963