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VOLUME LXVIII NUMBER 6

T H E

BOTANICAL GAZETTE
DECEMBER 1919

RELATIVE TRANSPIRATION OF CONIFEROUS AND

BROAD-LEAVED TREES IN AUTUMN AND WINTER'
J. E. WEAVER AND A. MOGENSEN

(WITH EIGHTEEN FIGURES)

Introduction
Transpiration has beenofspecialinterestto manyinvestigators
fora long time. At firstit was consideredwithoutreference to
environmental factors,but later,as moreobservations weremade
and thesefactors werenotedto havea markedeffect uponthewater
loss, they were taken into consideration.Many of the data
assembledhave been limitedto plantsduringthe growingseason,
so that it has seemedprofitable to obtainnot only quantitative
dataonwinterlosses,butalsoa comparison oftherelativetranspira-
tionof conifers and broad-leavedtreesin summerand winter.
Variousmethodshave been devisedfordetermining transpira-
tion,fromthecutshootpotometer, whichusuallygiveslossesquite
too low whencomparedwithrootedplants(6, I6), to the cobalt
chloridemethodofSTAHL (i7), recently improvedby LIVINGSTON
and SHREVE (I3). With few notable exceptions,such as the
methodusedby ILJIN(II), whoworkedon xerophytes and meso-
phytesin the field,the former methodhas been used largelyfor
laboratorymeasurements, while the latter,althoughespecially
devisedforfielduse, does not takeintoaccounttheenvironmental
I Contributionfromthe Department of Botany, Universityof Nebraska, new
series,no. 29.
393
394 BOTANICAL GAZETTE [DECEMBER

factorsof the habitat under which the plants are growingexcept

as measured by the internal transpiringpower of the plant, nor
does it give a recordof the continuoustranspirationlosses.
Unquestionablythe most reliabledata have been those obtained
by the potometer,where the entiresealed containerwas weighed
withthe wholeplant intact. A survey of the literature,however,
revealsratherscant data on the transpiringpoweroftrees,especially
coniferoustrees,regardlessof the method employed.
That evergreentrees are constantlysupplied with water, even
in winter,was firstobservedby HALES (8), and later by DUHAMEL
(i8), and others.
(4), TREvIRANUS
In i86o HARTIG(IO) made some investigationson transpiration
losses with Picea, a meterhigh, in milderwinter,and found that
the plant lost from about IOO-I25 gm. of water a day. These
figures,however,are oflittlevalue so faras calculatingthe intensity
of transpirationis concerned,since he gave neitherarea nor weight
of the transpiringpart.
BURGERSTEIN(3) in i875 indicatedthe relationof transpiration
to lower temperatures,and showed that cut branches of Taxus
baccatatranspiredin an hour, at - 2? C., 0. 288 per cent, and at
- 107O C., O.oI9 per cent of theirfreshweight.
That transpirationmay take place quite rapidly at ratherlow
temperatures has been shown by WIESNER and PACHER (22).
Twigs of Aesculusand Quercus,for example, lost 0.32 and 0.25
per cent respectivelyof their weight in 24 hours at -3 .50 to
-I . 5? C., and o. I99 andO.I92per cent at-5 .50 C. to-13 .00 C.
BEACH and ALLEN (I) found a loss of from4 to 9 per cent of
water in apple twigs during a single week in January, with a
minimumtemperatureof -26.o' C. They found also in general
that the hardiestvarieties were the most resistantto water loss.
According to WARMING(20, P. 310), coniferoustrees exhale
much less water vapor than dicotyledonoustrees, due to their
xerophyticnature.
KUSANO (I2) has given convincingquantitative data on the
transpirationof evergreentrees indigenous to Japan. He found
that evergreentrees transpiredin winter an average quantity of
at least o .48 gm.per sq. dm. per day (withthe exceptionofconifers),
I919] WEAVER &a MOGENSEN-TRANSPIRATION 395

or i6.58 gm. per ioo gm. of fresh weight in foliage trees, and
8.i8 gm. in conifers. He also noted that the time of minimum
transpirationagreeswith that of the minimumtemperature,which
occurred at the end of January. He states furtherthat the
differencein the amount of transpirationof differentspecies of
evergreentrees becomes smallest at the time of minimumtran-
spiration; and a change in the externalconditions,especially in
temperature,does not necessarilyproduce a correspondingchange
in transpirationin differentspecies. In average cases the amount
of water transpiredby foliage evergreentrees is one and a half or
two times greater than that transpiredby conifersif we reduce
the amount eitherto the freshweightor to the dry weight of the
transpiringpart.
VON H6HNEL (ig) estimated that a birch tree, with about
200,000 leaves and standing perfectlyfree,would evaporate 400
liters of water on a hot dry day. He also has calculated that
during the period of vegetation the beech requires 75 liters, and
the pine only 7 liters for every ioo gm. of leaf substance. The
same writergives us the followingtable on the relative amount
of water transpiredfromJune i to November 30 per ioo gm. dry
weightof leaf:
Birch . . . . 67. Oak . . . . 28.3
Lime . . . 6i .5 Red spruce . . 5.8
Beech . . . . 56.6 Whitepine . . 5.8
Maple . . . . 46.2 Silverfir . . . 4.4
Elm . . . . 40.7 Austrianpine . 3.2

Experimentson the transpirationof seedlingsof Acer sacchari-

numin prairieand shrubthickethave been carriedout by WEAVER
and THIEL (2I). Trees placed in the latter habitat lost only 30
per cent as much waterper sq. dm. as those in the prairie. Similar
experimentswith Quercus macrocarpa gave comparable results.
The high transpirationlosses in the prairie help to explain the
absence of trees fromsuch localities.
BERGEN (2) compared the transpirationrates of a number of
broad-leaved evergreens,including Olea europaea, Quercus h1ex,
and Pistacia Lentiscus,with those of equal leaf surfacesof Ulmus
campestrisand Pisum sativum. He found that the losses in the
396 BOTANICAL GAZETTE [DECEMBER

formergroup were only about 25 per cent less than in the latter,
and concluded"that xerophyticleaf structureis not always incom-
patible with abundant transpiration,but sometimesexists only for
use in emergenciesto protect the plant from injurious loss of
water."
HANSON(9) has shown that the major water losses fromthe
crowns of isolated trees of Ulmus americana, Acer saccharinum,
and Fraxinus lanceolata occur fromthe peripheralbranches, and
less than one-sixth from equal areas of leaf surface on shaded
branches.
The presentinvestigationwas undertakenwitha double purpose
of obtaining data on the relative losses in summerand winterof
conifersand broad-leaves, and also to make a beginningon the
problem of winterkillingof trees and shrubs. This latter project
is not included in thispaper.
Methods
In the spring of i9i8 seedling conifers of Pinus ponderosa
Dougl. and Pinus Banksiana Lamb. were obtained fromthe forest
nurseryat Halsey, Nebraska; while those of Abies grandisLindl.,
Pinus Murrayana Balf., Picea Engelmanni (Parry) Engelm., and
Pseudotsugamucronata(Raf.) Sudw. were securedfromthe national
forestsof northernIdaho. These seedlings,varying fromtwo to
fouryears in age, were potted duringMay in 5-, 7-,and 8-inchpots
respectively,accordingto the size of the plants and the demands of
the root systems,and in soil consistingof two parts of rich garden
loam and one part of sand, thoroughlymixed and screenedthrough
a one-fourth-inch-mesh sieve. The pots were placed on the lawn
near the greenhouseuntil needed in late summer,and were thor-
oughly watered every day and sometimestwice a day duringthe
driest periods. A few trees, mostly white fir and Engelmann
spruce, died. These, with numerous weaker individuals of other
species,werediscarded,and onlythe verybest plants,whichshowed
the most flourishingconditionof growth,were used in the experi-
ments; in fact, only about half of the original stock was thus
selected and used.
Galvanized iron containerswith flatbottomsand straightwalls
were used in the transpirationwork,the size varyingaccordingto
I9191 WEAVER &- MOGENSEN-TRANSPIRATION 397

the demandsof the root system. Thus, while the 2-year-old

Engelmannsprucerequiredcontainersonly 6.5 inchesdeep by
3.5 incheswide (thesewere the smallestused), the 3-year-old
yellowpinesweregrownduringthe experiments in containers5 .5
inchesin diameterand I4 inchesdeep. Such containers are very
desirable,sincetheycombinelightness withappropriate shape for
usingthe minimum amountofsoil.
In September, whenthe treesweretransplanted fromthepots
to the galvanizediron containers, theywere handledin such a
way as to scarcelydisturbthe root systems. A layerof coarse
gravel0.25 inchesdeep was placed in the bottomof each con-
tainer,to thesideofwhich,extending fromthebottomto thetop,
a heavy glass tube 5 mm. in diameterwas fastenedwithsealing
wax. The soil in the flowerpots, havingbeen well watered24
hourspreviousto transplanting, and the waterallowedto drain
through thebottomofthepot,wasofsucha texturethatthewhole
contentscouldeasilybe removedby inverting thepot and jarring
the edgewhileholdingthesoilsurfaceintactwitha pieceofcloth.
This core of soil, containingthe root systempracticallyundis-
turbed,was placed in the new container. In some cases it was
necessary to trimawaya partofthetop oftheconicalcore,butthe
part removedwas alwaysfreefromroots. Previouslythe sheet
metalcontainerhad been filledto such a depthwithsoil of the
samecomposition as that of the core,that the plant,whenput
in place,wouldbe at a properheightin relationto the top ofthe
pot. Soil samplesformoisturecontentdeterminations weretaken
at this time.. Any spaces in the new containerswere carefully
filledwithsoilwhichwas properly compacted. Severalplantshad
theirrootsystemsmoreor less disturbedin repotting, and these
werediscarded. Finally,thepotsweresealed.
The sealsconsistedofpetrolatum mixedwithparaffin;thelatter
had a meltingpointof about 50?C. Variousmixtures wereused,
from8o per centparaffin and 20 per centpetrolatum(by weight)
at thebeginning ofthe experiment, whenthe weatherwas hot,to
25 per centparaffin and 75 per centpetrolatumin midwinter.A
very satisfactoryseal forwinterweatherconsistsof the latter
mixture pouredon thesurfaceofthesoilwhileveryhot,and then
398 BOTANICAL GAZETTE [DECEMBER

coveredwitha less plasticseal of about equal parts of the two

ingredients.Littlediflfcultywas experienced in keepinga perfect
seal intact.
The broad-leavedtrees,Acersaccharinum L., Ulmusamericana
L., and Quercusmacrocarpa Michx.,werehandledin identically the
same manneras the preceding, exceptthattheyweregrownfrom
seed sowedin flatsin the greenhouse, but the plantsweretrans-
ferred to pots out ofdoorsin June.
In orderto insureuniform soil temperature conditionsforall
plantsconcerned, andchangesoftemperature similarto thoseunder

withcollarsto insuretemperature
oftreesin containers,
FIG. i.-Arrangement
withthoseofplantsgrowing
changescomparable undernaturalconditions.

naturalconditions, the containers(approximately ioo in number)

were placed in tin cylindricalcollars,each collar being slightly
largerthan its respectivecontainer. These were in rowsabout
6-io inchesapartand completely surrounded by soil. The top of
each container and collarwas coveredwitha heavywoolenblanket,
slitto accommodate thestem,and thisin turnwitha pieceofthick
muslinwaterproofed withthehotwax seal mixture.
by infiltration
The edgesespecially,and also thewholecloth,as wellas the sur-
rounding soil,werecoveredwitha thinlayerofsand. To prevent
thedrifting ofsnowamongthetreesthusplantedon theUniversity
campus,the collarsand inclosedcontainers wereset on the bare
soil surface,and the well-tamped soil filledin betweenthemwas
heldin place by a boardframeI2 incheshigh(fig.i.)
I919] WEAVER &aMOGENSEN-TRANSPIRATION 399

Continuousrecordsofthehumidity oftheair and thetempera-

tureofbothair and soilwereobtainedby meansofFriez'shygro-
thermographs.The last was fora soil depth of 6 inchesamong
the containers.A thermometer placed at a similardepthin one
ofthemediumsizedcontainers,andprotected fromexternalatmos-
phericconditionsby meansof a felt-linedbrass case fittedwith
a cap, gave readingsverysimilarto thoseof the soil,as may be
seenin tableI.
TABLE I
SOIL TEMPERATURE AT A DEPTH OF 6 INCHES INSIDE A
CONTAINER, AND AT A SIMILAR DEPTH
OUTSIDE IN THE SOIL

Date Temperature Temperature

among roots outside

DecemberI8 A.M..........
I.50C. . I .50C.
I8 P..2.0 2.0
3I -.4........... -4.0 -4.0
January I . -2.5 -2.5
2 . 0.0 0.0
3 .-.... .5 I.0
I4.............. -75 -8.o

Whilethefactordata willbe discussedin connection withthe

transpiration graphs,it maybe said herethatat no timewas the
soil in the containersfrozensolidlyto a depthgreaterthan2 .5-
3.0 inches,a pointextending notfarintotherootzone.
Transpirationlosses were determinedby weighingthe con-
tainers. A large long-armed Troemnerbalance was used which
was sensitiveto 0 . 5 gm.underthemaximumload ofabout 8-9 kg.
imposedupon it. In makingthe weighingsthe containerswere
transferred fromtheirplaceoutofdoorsintoa weighingroomcooled
to nearly a similar temperature,for,as shownby WINKLER(23),
leavesofevergreens and twigsofothertreescan endurefromfour
to six timesas muchcold if the changesare gradualas if they
are sudden. Unless the water loss was ratherinsignificant as
comparedwiththeamountofsoilconcerned, thepracticefollowed
was to replacethe loss at each weighingby addingthe proper
amountofwaterfroma burettethrough thetemporarily uncorked
glass tube in the side of the container. Thus the soil moisture
was keptat an almostuniform conditionthroughout.
400 BOTANICAL GAZETTE [DECEMBER

At leasttwoorthreetimes.during thecourseoftheexperiments,
whichextendedfromSeptemberI9I7 to May i9i8, the seal was
brokenand the soil thoroughly aeratedby meansof an aspirator.
An examination ofcontainers discardedfromtimeto timebecause
of accidentsto the aerial parts of the plantsshowedthat at all
timesthesoilwas sweetand in goodcondition. Such an examina-
tionalso revealedthatthe rootsofbothdeciduousand coniferous
treeshad penetrated somewhatintothenewsoil. As pointedout
by MACDOUGAL (14), therootgrowthofbroad-leavescorresponds
to the warmperiodsduringwhichabsorptionis active,whileany

old yellowpines; photographed

FIG. 2.-Battery Of3-year February28, igi8;
noneofneedlesfallen.

possiblerestingperiodin summeris deemeddue to scarcityof

waterand not to any inherenttendencyof the plant towarda
in growth.
periodicity
Pinus Ponderosa
Eight 3-year-old seedlings of yellow pine were repotted, as
alreadydescribed, 5 .5 inchesin diameter
intothemetalcontainers
and 14 inchesdeep,and thefirst weremadeon September
weighings
24. Thisbatteryis shownin fig.2. The totalleafarea was deter-
minedforseveraloftheplantsthefollowing springand at theend
oftheexperiment.Thisrangedfroma minimum Of2.905 sq. di.
in plant no. 7, to a maximumof 6.428 sq. di. in plant no. i
19I9] WEAVER & MOGENSEN-TRANSPIRATION 40I

(fig. 2). The areas were determinedby removingthe leaves from

the plants,measuringthe lengthof the leaf fascicleand the average
diameterof the flat faces, as well as the diameter of the cylinder
formedby the two or threeleaves in the fasciclewhen the flatfaces
were appressed against each other. From these data the actual
surface area was calculated. Practically no needles were shed
duringthe winter.
Fig. 3 shows the total average daily losses of six yellow pines
fromSeptember24 to January i. An examinationof these graphs
reveals a strikingsimilarity. The highest losses are from tree

(kt3- Ocf5- -
Oct./6- t.26- tNoV9-
OU 0\, /6 . Urv. XL J"

5y.
16C

FIG. 3.-Average daily losses in grams from six 3-year-oldyellow pines from
September24 to JanuaryI; heavy line representsmean temperatureforthe several
periods.

no. I with the greatestleaf area (6.428 sq. dm.), while the lowest
losses are plotted fromdata obtained fromtree no. 2, which had
an area of only 4. I sq. dm. Comparative losses per unit area are
given elsewhere. Data from the other two pines were omitted
in this figureforthe sake of clearness. A generalrelationbetween
temperatureand transpirationwas clearly evident. The relation
to humiditywas not so apparent.
The entireperiodfromSeptember24 to October i6 is character-
ized by relativelyhigh transpirationlosses, afterwhich there is a
decided fallingoff. On October ii the stomata were found to be
closed. The midwintertranspirationlosses are exceedinglysmall.
Weighingsmade on February 7 and aftera periodofprolongedcold
 402 BOTANICAL GAZETTE [DECEMBER

weather(the mean monthlytemperature of Januarybeingonly

I2.60 F.) gave total maximumlossesforthe entireperiodof 37
daysofonly2 .5 gm. The averagetotalloss duringthistimewas
about i gm. It is surprisinghow an area of 3-6 sq. dm. of leaf
surfacecan be exposedwith such minimumlosses. The daily
losses,compared withthosefromthesameplantsduringtheperiod
September 24 to Octoberii, are onlyl/25I as great. The average
lossesduringsucceedingintervalsare shownin tableII.

TABLE II
TOTAL AMOUNT (IN GM.) OF WATER TRANSPIRED BY 3-YEAR-OLD YELLOW PINES FROM
JANUARY I TO MAY 2, i9i8

Januaryi- February7- MarchI3-26 March26- March26-

Plant February7 MarchI3 ( days) April29 May 2
(37 days) (34 days) (34 days) (37 days)

I............ 2.0 13.7 8.8 7.3 ............

2 ........ 0. 5 ............ 12.2 i6.5 ............
3 ........ 2.5 9.5 11.0 13.5 ............
4 ........ I5. 8.o 14.5 ............ 69.o
5 ........ I5. 79 i6.i ............ 70.0
6......... 0.5 10.2 13.8 . ........... 26.o
7-.. ........ 0 -5- 4 -5 ............ ............ . .............
... .
8. .......... O 5 13.1 23.9 . .. 91I.4

A generalbut slowriseduringthe coldmonthofAprilmaybe

noted,witha sharpincreasefollowing themilderweatherin May.
The transpiringarea throughout constant,sincelittle
is practically
growth occurredbeforeMay i. On 3
April the stomatawerefound
to be open.
Initialweighingsfroma batteryoffive2-year-old yellowpines,
each in a 4X9-inchcontainer, wereobtainedon Octoberi8. Two
oftheseplantsareshowninfig.I 2. Table III givesthetotallosses
duringthe severalperiods.
On Decemberi9 the leaves werefallingso badly that all but
two plants werediscarded,but the following springall put out
new leaves.
The exceedinglylow waterlossesduringthe periodDecember
I9 to March I3 correspond withthosefoundfortheotherbattery
of olderyellowpines; in fact,thiswas foundto be the case with
all oftheconifers.Table III showsan increasein the transpiring
I919] WEAVER P MOGENSEN-TRANSPIRA TION 403

poweras springadvancedsimilarto that of the older pines in

the otherbattery. These plantsweregrowingin a soil withan
averagewatercontentof I5.8 per cent(i8.2 per centmaximum,

TABLE III
TOTAL TRANSPIRATION LOSSES (IN GM.) OF 2-YEAR-OLD YELLOW PINES FROM
OCTOBER i8 TO MAY 2

October i8- November i7- December i9- March I3- April 2-

Plant November i7 December i9 March I3 April 2 May 2
(30 days) (32 days) (84 days) (20 days) (30 days)

I.. ....... 44.0 12.0 ............ ............ ............

2.. ....... 48.0 14.1 ............ ............ ............
3.... ....... 42.0 4.0 ............ ............ ............
4.. ....... 17.66. I 4.3 8.2 13.6
5......... 22.5 6.5 3.4 6.3 12.5

I4.6 per centminimum), and an availablewatercontentvarying

fromI3 .5 to IO per cent.
PinusBanksiana
A battery ofeight3-year-old jack pinesin containers 4.5 inches
in diameterand IO inchesdeep was sealed and weighedon Sep-
tember26. Thesetreesweregrowing in a soilwithan actualwater
contentranging from 8.6 to I5.I per cent,of whichonly4 and
Io.3 per centrespectively wereavailableforgrowth.
The leaf area, calculatedin a mannersimilarto that already
described fortheyellowpines,rangedfrom2 . I4I to 4.470 sq. dm.
The plantsremainedin good conditionthroughout thewinterand
showedvigorousgrowthin the spring. In additionto the usual
brownishcolorof the leaves in winter,however,the tipsof many
of the leaves died duringJanuaryand February. Practicallyall
the leaves remainedon the plants throughout the experiment.
Fig. 4 showsthisbatteryas it appearedon February28.
The transpiration lossesas determined forthe severalperiods
are plottedin fig.5. A glanceat thesegraphsshowstwo things
whichare at once apparent. First,the generalparallelismof the
linesthroughout (exceptfromOctoberI7 to 26, to be considered
later); that is, the plantwhichgave the highestor lowestlosses
duringtheearlyperiodscontinued thisbehaviorthroughout.The
404 BOTANICAL GAZETTE [DECEMBER.

actuallossesaregenerally
correlatedwiththeleafarea; forexample,
thegreatestlossesarefromplantno. 7 withan areaof4.47 sq. dm.,
whilethosein graphno. 2 are froma plantwithan area of only
2.I4 sq. dm. Secondly,theremay be noteda riseor fallin the
graphswhichcorresponds in generalwith temperaturechanges.
AfterOctoberI 7 therewas a gradualbutmarkedfallingoffinwater

April, when growthwas resumed. These data are shown in

tableIV.
TABLE IV
TOTAL TRANSPIRATION LOSSES (IN GM.) FROM 3-YEAR-OLD JACK PINES DURING THE
SEVERAL INTERVALS FROM DECEMBER I9 TO APRIL 29

DecemberI9- Decemberig- March26-

Plant February7 March26 April29
(5o days) (97 days) (34 days)

I .......... .............. 39 5 96.4

2............. .............. 14.0 36.5
38 ............. .............. 8. o48-5
4 ............. .............. 24.5 89.9
5 ............. .............. 20.2 56-5
6 ............. 30 February7-
March26
(47 days) ..............
7............. 2.0 I2.0 ..............
8 ............. 3-5 30.8 90.0
I9I9] WEAVER P MOGENSEN-TRANSPIRATION 405

The midwinter losses,actualand relative,arejust as markedas

they were in the case of the yellowpines. During the period
Decemberi9 to February7 thetotalaveragelosswas only2 .8 gm.,
an amountwhich,comparedonthedailybasis,is onlyi/i69 ofthat
lost duringtheautumnperiod(September26 to OctoberI 7).

Abiesgrandis
Two batteriesofwhitefirswereused. One consistedof eight
2-year-oldseedlingsin containers3 .5 inchesin diameterby 6.5
inchesdeep,and the otherof eight4-year-oldtreesin containers
3 .5 inchesin diameterand 9 inchesdeep.

p.2-Ocea-&-t10 0&17- OcZ26- o.

FIG. 5. Averagedailylossesin gramsfromeight3-year-oldjack pines; September

26 to December i9, I9I7.

The younger plantsweregrowing in soilwithan availablewater

contentrangingfrom6 to 9.4 per cent. All werewinterkilled.
As thisdid notoccuruntilJanuaryor February,thetranspiration
lossesup to thattimeare reliable. AboutJanuaryI5 the leaves
beganto dropoffbadly,but the leafarea of severalof the plants
was determined withoutremovingtheleaves,andbeforedefoliation
had begun. Thiswas accomplished by determining thenumberof
leaves, their lengths,and average diameters. Three of these
seedlings had leaf areas of 0.0972, o.i64I, and 0.0970 sq. dm.
respectively.These plants are shownin fig. i i. This picture
was takenlate in Februaryaftermanyof the leaves had fallen.
Because of an accidentto the top ofone tree,it was discarded.
406 BOTANICAL GAZETTE [DECEMBER

The transpirationlosses fromSeptember27 to DecemberI2

fromthesevenremaining plantsare shownin fig.6. The average
total loss duringthe periodof DecemberI2 to Februaryi i was
only0.7 gm.,the maximumand minimum lossesbeing I .0 and
o .3 gm. respectively.This dailywinterloss comparedwiththat
of earlyfall(September27 to Octoberio) is only I/55 as great.
It mustbe noted,however, thatit was duringthisperiodthatthe
seedlingswerewinterkilledand may have lost morewaterthan

p127- Ok1/O-
Zi/
tYO1/6-
0123. 1T6
0d23-
2

FIG. 6.-Average daily losses in grams fromseven 2-year-oldwhite firsfrom

September27 to December I2, I9I7.

normally. On Februaryi i the leaveswerebrownand theplants

appeareddead.
The 4-year-oldfirs,althoughgrowingundermore favorable
moisture conditions (from5.3 to I3 .7 percentavailablemoisture),
also succumbedto thedrycoldwinter. On Februaryi i theleaves
wereall brownand dead. The transpiration lossesto Decemberi9
are reliable,however, forup to thistimeall theplantswerein good
condition. The leaves did not dropbadly,even afterdrying,as
is shownin fig.7, a photograph whichwas takenon February28.
Leaf areasforplants5 and 6 werecalculatedas fortheotherwhite
firs. These leafareas werei.o65 and o.638 sq. din. respectively.
I919] WEAVER & MOGENSEN-TRANSPIRATION 407

Sinceweighingsofbothsetsoffirsweretakenon thesamedays,
thegraphsin fig.6 maybe compareddirectlywiththosein fig.8.

FIG. 7.-Battery of 4-year-old

whitefirs;photographed
February28, I9I8,
aftersomeofleaveshad fallen.

FIG. 8.-Averagedailylossesin gramsfrombatteryof seven4-year-old

white
firs; September28 to December i9, I917.

A markedirregularityin the courseof the otherwisegenerally

parallelgraphsmay be seen duringthe periodof October23 to
Novemberi6.
408 BOTANICAL GAZETTE [DECEMBER

Attentionhas alreadybeen calledto thisphenomenon in case

ofboththeyellowand jack pines,and figs.9 and io showthesame
forthespruceand Douglasfir. Althoughthisphenom-
occurrence
enonwas notexaminedcarefully at thetime,forit was notknown

ept28- OcA
/J- Oct23- A~oM9

FIG. 9.-Average daily losses in gramsfrombatteryof eight3-year-oldspruces;

September28 to December i3, I9I7.

w27- OcdlO- ob/23-

FIG. iO.-Average daily losses in gramsfrombatteryof three3-year-oldDouglas

firs; September27 to December i9, I9I7.

to occuruntilafterthefinalweighingsonNovemberi6 wereplotted,
the irregularity shownby the
is probablydue to the individuality
severalplantsin therapidityofpermanent closureofthestomata,
and a generalslowingdownofthevitalactivitiesas thetemperature
decreased.
I919] WEAVER & MOGENSEN-TRANSPIRATION 409

Picea Engelmanni
This batteryconsistedof eight3-year-oldEngelmannspruce
seedlingsgrownin containers3.5 inchesin diameterby 6.5 inches
in depth,and in soilwithavailablemoisturerangingfromII.3 to
15 . I per cent.
Althoughthe leaves droppedoffbadly duringlate December
and in January, noneoftheplantsdied. All put fortha vigorous
growthofnewleavesduringApriland May ofthefollowing spring.
Threeofthesetrees,photographed inFebruary, areshowninfig.I I.
... . . .. . .... . . . . ... . ..

. ... .

.*

FIGV iiThree 3-year-old three2-yearold white

Engehannspruces, firsand
allmore
oneDouglasfir, orlessdefoliated; takenFebruary
photograph 28, 198

The leafareasofthreeplants,calculatedat thetimetheleaves

had just begunto fall,were2.78, 3.91, and 5.87 sq. din. respec-
tively. These areas weredetermined whilethe leaves werestill
intactby countingtheirtotal number,measuringthe lengthand
averagediameterofeach leaf,and thencalculatingthearea of the
foursides. The graphsgivingthe transpiration lossesare shown
in fig.9. A comparison ofthewaterlossesfromtheseplantswith
thoseof otherconifersshowsa remarkable similarity.

Pseudotsugamucronata
Because of highmortality
amongthe Douglas firsduringthe
reestablishmentin pots in early summer, only three 3-year-old
seedlings were available for experimental work in September.
Thesewereplacedin containers 3,. inchesin diameter
by 8 inches
4IO BOTANICAL GAZETTE [DECEMBER

in depth,sealedand weighedon September27. The watercontent

of the soil in the three containerswas I3.8, I5, and i8 per cent
was 4.7.
respectively.The wiltingcoefficient Oneoftheseplants
is shownin fig.II.
The leaf area of plant no. I was o .3062 sq. dm. This was
determined by considering the leaves as havingtwo flatsurfaces
and multiplying thelengthofeachby its averagediameter.
Transpiration lossesare shownfromSeptember27 to December
i9 in fig.io. During theperiodDecemberi9 to FebruaryII, the
total losses of two plants (the thirdhaving accidentallybeen

FiG. 12.-Batteryof five3-year-old lodgepolepinesand two 2-yearold yellow

pines;photographed February28, 1r9i8.

brokenoff)wereo 4 and i . i gin. respectively On the basis of

the average daily loss, this is a mere fraction(only I/I46) of that
duringtheautumnperiodSeptember27 to Octoberio. In March
theplantswerediscardedbecauseofthefallingleaves; in fact,one
finallydied,but the othersshowedrenewedgrowthin the spring.
Pinus Murrayana
Still anotherbatteryof needle-leaved of six
plants,consisting
3-year-old lodge pole pines, was experimentedupon during the
periodbeginningOctoberI8 These plants,grownin containers
4 inchesin diameterand 9 inchesin depth,are shownin fig.
si2.
The watercontentof the soil rangedfromi6.4 to 2I .8 per cent.
I919] WEAVER & MOGENSEN-TRANSPIRATION 4II

All of the plants died in February,although the needles persisted

fora long time.
The transpirationlosses shown in table V are not greatly
differentin their amount fromthose of the other conifers. Like-
wise they show a gradual decrease as winter approaches, with
minimum midwinter losses. All losses recorded are reliable
because no readingswere taken after the plants showed signs of
deterioration.
TABLE V
TOTAL LOSSES (IN GM.) FROM A BATTERY OF SIX LODGE POLE PINES FROM OCTOBER i8
TO FEBRUARY II

October i8- October 26- November 17- January 4-

Plant October 26 November I7 January 4 February ii
(8 days) (22 days) (48 days) (38 days)

I................ 4-.5 9.5 3.9 0.7

2................ I3.4 30.4 6.o 0.7
3................ 2.9 I5.5 5-5 I 6
4................ 5.3 I3-5 2.6 0.7
5................ 7- 5 I5-5 7.8 I.I
6............. II.5 37-7 II.3 I.0

Ulmus americana
Initial weighingsof a batteryof I2 white elm trees were made
on September 20. These plants were in containers 5 .5 inches in
diameter and 8.5 inches deep. A photograph taken on May i
just when the plants were leafingout is shown in fig. 13.
The leaf areas were determined,as in the case of the other
dicotyledons,by means of solio leaf prints. These were made,
of course,withoutremovingthe leaves fromthe stems. The areas
of the plants whose transpirationlosses are shown in fig. 14 were
as follows:
No. 6 . . . IO.28 Sq. dm. No.IO . . 4.I5sq.dm.
7 * 8.57 II * * * 4.52
8 . 6.56 12 . . . 5. 26
9 * 8.97

For the sake of clarity,the losses fromthe other plants were

not recordedin fig. 14. In all cases theywere verysimilar. With
the exceptionof plant no. 12, there is a close agreementbetween
412 BOTANICAL GAZETTE IDECEMBER

puttingout newleaves; photographed

FiGo13.Battery ofwhiteelmis May i

epi'O-ept4- ep~28- 64/5- Ocflf

/6

elmtrees; Sep-
FIo. 14.-Averagedailylossesin gramsfromseveni-year-old
tember20 to October26, 1917.
I919] WEAVER &eMOGENSEN-TRANSPIRATION 413

the early autumnlosses (September20 to October i i) and the

initialleafarea. Alsoa comparison ofthegraphofmeantempera-
tureforthe severalperiods(fig.I 5), withthe graphsshowingthe
marchoftranspiration,showsthattheyhavea striking resemblance.
The meantemperature wasobtainedfromthethermograph records,
and is givenon a time-humidity basis. The meansfortheseveral
periodsrespectively were obtainedby drawinga horizontalline
throughthe weeklyrecordsheetin such a mannerthat the total
area includedby the graphabove thisline was equal to the total
area below the line. The areas weredetermined by the aid of a
planimeter. In this interpretation both temperatureand time
factorsare takenintoconsideration.Temperature, indeed,seems
factor,forno suchcorrelation
to be the controlling betweentran-
spirationand humidity (herethe graphis inverted)is discernible
afterOctober5; that is, transpirationdroppedoffat this time,
following thedescending temperature graphinspiteofa decreasing
humidity.
These two factors,however,togetherwithwind velocity,are
well summedup in evaporimeter readings. These wereobtained
by Livingston'sporouscup atmometers and showan increaseor
decreasevery similarto the transpirationgraphs. They are as
follows:
September
20-24 . 25. I cc. averagedailyloss
September
24-28 . ii .8 cc.
28-October
September 6 . 2.2 cc.

Readingswerediscontinuedon Octoberi i because of freezing

weatherat night. Unfortunately no instrument has yet been
devisedfor measuring
satisfactorily winter evaporation losses.
in thegraphs,especiallyin the fourthinterval,
The irregularity
is due to the non-uniform dryingout of the leaves. Up to October
5 the leaves of all the dicotyledonousplants were greenand appar-
ently functioningnormally. By October ii a few of the older
basal leaves were beginningto turn brown. While some dropped
at once, many held on afterthey were dry. This irregularityin
defoliationdefeated an attempt to base the daily losses upon the
actual average leaf area duringany givenperiod,althoughan exact
4I4 BOTANICAL GAZETTE [DECEMBER

recordwas keptofthetimeofthefallofeachleaf,together withits

area.
The transpiration graphs show a general fallingoff after
Octoberii, whichis notunlikethatoftheconifers;onlyherethe
transpirationis froma constantlydecreasingarea (all the leaves
had fallenor remainingfragments wereremovedby October26),
whilein the conifersthe area remainsconstant. In table VI are
giventhetotallosses(whichare too small to plot on an average
dailybasis) fromOctober26 to May i.

TABLE VI
TOTAL LOSSES (IN GM.) FROM I 2 ELM TREES FROM OCTOBER 26 TO MAY I

October26- November23- January 4- April 24-

Plant November9 January4 April24 May I
(14 days) (42 days) (iio days) (7 days)

I. .............. .......... I.0 7.4 2.2

2 . ............. I'.0 5.6 3.2
3........... 6.9 I.0 I2.3 12.7
4 ........... 2.8 I. 0 6.6 2.7
6........... 33 I.0 II .3 9 7
7---............ 3.4 1.5 Io. 8 1.3
8 ........... 2.0 1.0 5.8 3.2
9........... 5-7 I.5 9-3 37
Io .............. I. 9 0. 5 7-0 5.8
II ............ 0 .4 0.5 4 5 2.3
I2 ............. I1. 5 0.5 7.I 2.2

FromNovember9 to 23 the seals wereremovedand the soil

allowed to dry out. The loss was about I25-I75 gm. They were
thenresealed,and 250 cc. ofwaterwas added. On April24 some
oftheplants,as nos. 2, io, and ii, had buds swollenbut not yet
open,whileinothers,as nos. 3, 6, and 7, theleaveswereunfolding.
On May i nos. 3 and 6 had leaves fairlywellout, some of them
beingoverI . 5 cm.long,whilein others(as nos. i and 7) theleaves
werejust bursting fromthebuds.
The winterlossesare verysmall. Duringa periodof 42 days
(November23 to January4) theaveragelosswas onlyi gm.from
the bare stemswhichexposedan area of 0.28 to o.5o sq. dm.
Table IX gives the relativetranspiration losses of conifersand
broad-leavedtrees.
1919] WEAVER & MOGENSEN-TRANSPIRA TION 4I5

Acersaccharinum
Weighings ofa batteryofI2 softmaplesweremadeon thesame
dates as thosefortheelms. The containers wereofthesamesize,
Sepft2-Sevi,24- Sep*._28- Oct5- 0cM8
5t8. O'a. co
/8. I

(solidline)and meanhumidity
FIG. I 5.-Mean temperature line)during
(broken
severalintervalsfrom 20 to October
September :26, 1917.

qqa'*0-Sept24- SevtO-;8 06cts OcH//- 0c./

0C5.
54tS/Qi28 /Oc: /S-
/YOY

s 05 vtember20 to October26 1917

FIG. i6.-Average daily losses in gramsfromseven i-year-oldmaple trees; Sep-

tember20 to November 2, 1917.

andthebatteriesand themethodsofhandlingthemwereso similar

thatfurther
throughout willbe unnecessary.
description
416 BOTANICAL GAZETTE [DECEMBER

The leafareasrangedfromII .32 sq. dm. in plantno. 6, which

gave the highestlosses throughout(fig. i6), to 6.47 sq. dm. in
no. ii, wherethe transpirationlosses are amongthe lowest. A
good correlationexistedbetweenleaf area and the magnitudeof
the losses. To avoid confusing of lines,data from
a multiplicity
fiveofthemapleshave been omittedin thisfigure. The data of
thesefivetreesare verysimilarto thoseshown.

TABLE VII
TOTAL LOSSES (IN GM.) FROM I2 MAPLE TREES FROM NOVEMBER 2 TO MAY I

Plant
Pat November November 28-
ii January April24-
i Conditionon
2-I6*
(1 2 days) January
(34 days) April24i- May nMyi
on May
(113 days) (7 days)

2 ; 2. 2 o.8 2.9 3.6 Two buds un-

foldingleaves
6 2.5 o.8 4.5 2.2 Just showing
signs of life
8 .3.0 I. I ........... ........... ................
IO 1.
I7 0.3 3.6 70.I Leaves I-5 cm.
long
I2 ........ I. 8 0.5 2.3 I3.5 Two buds with
leaves 2 cm.
long
December 6-
JanuaryI
(26 days)
I .... . .. . i .8 .... ... . . . . .. . . .. . ... . .. . .. ... .. . . . .. .... .. .. ...
3 ....... 2.0 ........... 3.7 4.0 One bud swollen;
two others
partly open
but frozen
November2-6*
(14 days)
5 .1..... . I .5 0 .9 ........... ........... ................
7 ........ 2 .0 i .8 .......... .. ......... .. ..............
9 ........ I .5 I .0 ........... ........... ................
II 2.5 0O5 ...................... ................

* All leavesoff.

As in the case of the elms,a close correlationexistsbetween

transpiration, and temperature
evaporation, (fig.I5); in fact,the
generaltrendofthegraphsis strikinglysimilarto thoseoftheelms.
The same irregularity is shownduringthe periodof defoliation.
On October26 onlythebare stemsremained;thesehad a surface
area of0. 22-0.48 sq. dm.
FurtherlossesfromNovember2 to May i are givenin table
VII. The smalllossesare againevident,followedby a rapidrise
I919] WEAVER &r MOGENSEN-TRANSPIRA TION 4I7

accompanyingthe unfoldingof the buds. Six plants were winter

killed, and no losses were recordedforthemafterJanuaryI.

Quercus macrocarpa
A numberofbur oak seedlingsweregrownfromacornsobtained
froman Illinois nurseryin May I9I7. Seven of the lot werefinally
selected forexperimentalwork. These were repottedin containers
5 inches in diameterand 7 inches deep. They may be seen in the
foregroundin fig. I. The initial available water content of the
soil ranged from II .3 to I5.2 per cent, but the soil moisture was
reduced by evaporation from the surface of the several containers

TABLE VIII
AVERAGE DAILY LOSSES (IN GM.) OF BUR OAK SEEDLINGS FROM SEPTEMBER 24 TO
JANUARY 4

P
Plant September 24- October 3-1O October ro-r8 November I-I6* December 7-
October)( (8 days) January 4
(9d days) (7 days) (I5 days)
~~~~~~~~~~~~~(28
days)
I ........... 9. 0 7.0 2.1 0.25 1.0
2 ........... ........... . 11.0 4-9 0-55 1.3
3........ . 10.2 6.4 4-3 1.22 1.4
4 . .......... 15-5 9.9 I.I 0.26 i. 8
5........... 7.2 3.4 0.4 0.2I I.0
6. .......... 4-5 2.6 0.5 0.28 1.0
7........ 15.2 I2.9 8.o I.89 i .8

* At this time all the leaves had fallen, and the losses are not calculated on the average daily basis,
but on the total loss forthe entire period.

to a minimum of 4.4 and a maximum of 9.6 per cent respec-

tively from October I8 to November I. After this the seals were
again replaced and half of the containers brought back to the
original water content. The transpiration losses are shown in
table VIII.
The gradual falling off in the transpiration rate without any
increase, as in the case of the elms and maples, may be explained
by an examination of the dates, which show that the firstloss was
not recorded until a time (October 3) when all the trees show a
steady decrease. By November I all of the leaves had fallen
(or the dried leaves were removed). Following this the losses are
extremely small. The stem areas of the oaks were only 0 039-
o.o88 sq. dm.
4I8 BOTANICAL GAZETTE [DECEMBER

Relative transpirationrates
Thus far we have considered largely transpirationlosses of
differentindividuals of the batteries of the various species, with
only occasional referenceto comparativelosses of different species.
It may be well,therefore, to presentdata showingthe actual losses
of differentspecies based upon unit area. These data have been
summarizedin table IX. While the dates between readings are
not synchronousin all cases, theyare at least nearlyso, and fig.is,

TABLE IX
COMPARATIVE TRANSPIRATION LOSSES FROM UNIT AREAS OF SURFACE OF BROAD-
LEAVED AND EVERGREEN TREES

Species No. of Time Average. loss per

trees sq. din.perday

Acer saccharinum .9 September24-October 5 2.66

Ulmus americana............. 9 " 24- " 5 3.56
Quercus macrocarpa.......... 7 " 24- 3 5. i8
Picea Engelmanni............ 3 " 28- 10 4. i8
Abies grandis(2 years old)..... 3 " 27- 10 5.44
Abies grandis(3 years old)..... 2 " 28- II 4.76
Pinus ponderosa.............. 3 " 24- " 5 4.20
Pinus Banksiana.3 26- " O 4.80

which gives the temperatureand humidity,together with the

record fromthe atmometers,shows that the overlapping periods
are quite uniform.

Relative midsummertranspirationrates of broad-leaved trees

and conifers
During the followingsummer,while the writerswere carrying
on otherinvestigationsin the Rocky Mountains of the Pike's Peak
region of Colorado, opportunity was afforded to compare the
summertranspirationrates of different tree species.
Seedlings of white elm and soft maple grownfromseed at the
Alpine Laboratory near Manitou, Colorado, and at an altitude of
8ooo ft., together with mountain maple (Acer glabrum Torr.)
obtained from the forest, were the broad-leaved species used.
Yellow pine, Douglas fir,and Engelmann spruce were the conifers
employed. The pines were secured fromthe Fremont Experiment
Station; while the other trees, like the mountain maples, were
1919] WEAVER &' MOGENSEN-TRANSPIRA TION 4I9

transplantedinto appropriatesized containersdirectlyfromthe

forestfloor,withoutdisturbingthe rootsystem. Care was taken

.~~ ~~~~~~~~~~
~ ......

FIG. 17.-Battery of Douglas firsand Engelmann sprucesfromwhichsummer

transpirationlosses wereobtained.

~~~~~~~~~~~~~.........
._ _ .. ....

FIG. I78 Battery of yellow pines and mountain maples used for determining
relativesummertranspiration.

not to selectthesespeciesfromtoo denseshade. Some of these

trees are shown in figs.I7 and i 8.
420 BOTANICAL GAZETTE [DECEMBER

The extrasoil necessaryto fillthesecontainers was in all cases

a mixtureoftwopartsofgardenloam and one partoffinegravel.
of IO . 7 per cent,and was keptrather
It had a wiltingcoefficient
uniformly at a watercontentofabout 20 per cent. The methods
usedthroughout werethesameas thosealreadydescribed.Original
weighingswerenot taken until afterthe treeshad been in the
containerstwoweeks. At thistimerabbitsattackedthedeciduous
treesand injuredall but fiveofthem.
On July20 theremaining treesintheircontainers, I9 innumber,
wereweighedto the nearesthalfgram. As duringthe preceding
interval,theywerethenplacedin a rowin the soil and in sucha
positionthat all were shaded fora fewhoursin the afternoon.

TABLE X
AVERAGE TRANSPIRATION LOSSES PER UNIT AREA FROM
SEVERAL SPECIES OF TREE SEEDLINGS

Species
No. of Averageloss
plants persq. dm.

Pinus ponderosa............ 4 58. 2

Pseudotsugamucronata .. . 5 50-7
Picea Engelmanni.......... 5 58.3
Acer glabrum........ ...... 4 II2.5
Acer saccharinum. . . . . . I I39.3

At theend of26 days,on AugustI5, theywereagainweighed,the

leafareasdetermined, and thelossescalculatedas in thepreceding
experiments.Thesedata areshownin tableX.
These data show that the losses fromthe variousspeciesof
growingin the openbut underconditions
conifers, whereall were
shadedfora portionof the day, are verysimilar,and onlyabout
halfas greatas thoseofthebroad-leaved species. The midsummer
averagedailylossper squaredecimeter fromthe yellowpinesand
Engelmannspruceat theAlpineLaboratory, whencomparedwith
thatofthesamespeciesduringtheautumnat Lincoln,was found
to be onlyabout halfas greatat the former station. Because of
the limitednumberofplantsused,however,thesedata shouldbe
consideredindicativeratherthan conclusive.
I919] WEAVER & MOGENSEN-TRANSPIRATION 421

Conclusions
A consideration of the foregoing data leads us to some con-
clusionsquite the converseof statementsgenerallycurrentin
literature.Perhapsthemostimportant
ecological-physiological of
these are the factsshownto hold underthe conditionsof these
experiments;first,that broad-leavedtrees underlate summer
conditionshave no greaterand indeedoftena smallertranspiring
power,area forarea, thanconifers;and secondly,thatthewater
lossesof coniferous treesduringthe wintermonthsare relatively
nogreaterwiththe needles intactthanarethelossesfromdeciduous
treesafterthe leaveshave fallen.
The factthatall oftheplantsconcerned, including threespecies
and about 30 individualsof broad-leavedtrees,and six species
represented by about 70 individualsofconifers, gave resultswhich
withoutexceptionpointto theseconclusions, leaves littledoubt
as to the validityof the finds. These resultswereobtainedwith
all the plantsunderuniform conditionsof soil type and texture,
soil temperature, and identicalaerial environment.The only
variablefactorwas soilmoisture, and heretherangewas no greater
forthe one groupthanforthe other. Althoughthe experiments
wereundertaken near the end of the summer,fora periodof a
fewweeks,aftertheirbeginning thedeciduoustreeswereinexcellent
growingcondition,and the comparativelosses as here recorded
occurredduringthe earlierpartof thisperiod,whenthe weather
was similarto that of midsummer as regardstemperature and
humidity.
That winterlosses fromthe same leaves that transpiredso
freelythe precedingfall and again in the following springare so
smallis certainly
testimony oftheecologicalefficiencyofconiferous
leaf structurefor reducingwater losses. Whetherthis is due
entirelyto stomatalclosure,or, as seemsmoreprobable,is con-
nectedwithchemicalchangesin cell contentsas well,remainsto
be determined.Such workas that of MIYAKE (Is) on the food
makingofconiferous leaves in winterand EHLERS (5) on tempera-
tureis rapidlythrowing lightupon the winteractiv-
considerable
itiesof coniferoustrees.
422 BOTANICAL GAZETTE [DECEMBER

The gradualdeclineof the transpiring powerof broad-leaved

treesprecedingdefoliation
and duringtheformation oftheabsciss-
layer is what mightbe expected,but is heretofore, we believe,
unrecorded.Weather conditionsplay an importantpart in
hastening orretarding
thisprocess,and as a resultthedropping off
in the transpiration
graph. These graphsare surprisingly similar
in theirgeneraltrendto those of coniferswherethe leaves are
intact.
The springof i9i8 was unfavorableforrapid leafingout or
renewedgrowthoftrees. Consequently the increasedtranspiring
power,whichis in a directratiowiththe leafarea, occurredvery
gradually,and hereagainwasnotgreatlyunlikethatoftheneedle-
leaved trees. Undoubtedly underveryfavorableweathercondi-
tions,as occurduringsomesprings,the transpiring powermight
easilydoubleor trebleday by day fora ratherlong consecutive
period.
Summary
i. Autumntranspiration lossesfromconifers are just as great
as or evengreaterthanthosefrombroad-leaves.
2. The decrease inwaterlossesfrombroad-leaved treesresulting
fromdefoliation is gradual,and not greatlyunlikethe decrease
shownin thetranspiring powerof conifers.
3. Wintertranspiration losses fromconifersare only I/55-
I/25I as greatas thosein autumn.
4. The increasedlosses of broad-leavedtreesin springocca-
sionedby foliationarein proportion to theleafareasexposed,and
are closelycontrolled by weatherconditions, but in the mainare
similarto increasedlossesof conifers.
5. Winter transpirationlossesfromconifers are scarcelygreater
thanthosefromdefoliatedstemsofbroad-leavedtrees.

The writersare indebtedto Dr. R. J. POOL forhis kindly

interestand encouragement,and to Dr. L. J. BRIGGS forwilting
coefficient
determinations;also to Mr. J. HIGGINS, of Halsey,
Nebraska,and Mr. T. J. LARSEN, PriestRiver,Idaho, fortree
I9191 WEAVER & MOGENSEN-TRANSPIRATION 423

seedlings fromthe nurseries,and to ProfessorT. J. FITZPATRICK

for carefulreading of the manuscriptand proof.
UNIVERSITY OF NEBRASKA
LINCOLN, NEB.

LITERATURE CITED
I. BEACH, S. A., and ALLEN, F. W., Jr.,Hardinessin the apple as correlated
with structure and composition. Iowa Agric. Exp. Sta., Research Bull.
2I, p. i85. I9I5.
2. BERGEN, J. Y., Transpirationof sun leaves and shade leaves of Olea
europaea and other broad-leaved evergreens. BOT. GAZ. 38: 285-296.
I904.
3. BURGERSTEIN,A., Vber die Transpirationvon Taxus-Zweigenbei niederen
Temperaturen. Oesterr.Bot. Zeitschr.25: i875.
des bois. I:337. I764.
4. DUHAMEL, H. L., De l'exploitation
5. EHLERS, JOHN H., The temperature of leaves of Pinus in winter. Amer.
Jour. Bot. 2:32-70. I19I5.
6. FREEMAN, GEO. F., Method of determination of transpirationin plants.
BOT. GAZ. 46: ii8-I29. i908.
7. HABERLANDT, G., Physiological plant anatomy, p. I38. Trans. from 4th
German ed. by MONTAGUDRUMMOND. London. I19I4.
8. HALES, STEPHEN, Statik der Gewachse, p. 29. I748.
9. HANSON,H. C., Leaf structure as related to environment. Amer. Jour.
Bot. 4:533-560. I9I7.
IO. HARTIG, T., Vber die Bewegung des Saftes in den Holzpflanzen. Bot.
Zeit. I9:I7. i86i; and LehrbuchfurF6rsterI:252. I877.
II. ILJIN, V. S., Relation of transpiration to assimilationin steppe plants.
Jour.Ecology4:65-82. 1916.
I2. KUSANO,S., Transpirationof evergreentreesin winter. Jour. Coll. Sci.,
Imperial Univ. Tokyo I5: part 3. I9OI.
I3. LIVINGSTON,B. E., and SHREVE, EDITH B., Improvements in the method
power of plant surfacesby hygrometric
the transpiring
for determining
paper. Plant World Ig: 287-309. I9I6.
14. MACDOUGALL,W. B., The growth of forest tree roots. Amer. Jour.Bot.
3: 384-392. I9I6.
I5. MIYAKE, K., On the starch of evergreen leaves and its relation to photo-
synthesis during the winter. BOT. GAZ. 33:32I-340. 1902.
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