Surface Dust Impacts on Gas Exchange in Mojave Desert Shrubs

Author(s): M. Rasoul Sharifi, Arthur C. Gibson and Philip W. Rundel

Source: Journal of Applied Ecology, Vol. 34, No. 4 (Aug., 1997), pp. 837-846
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/2405275
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Journalof
AppliedEcology
Surfacedustimpactson gas exchangein Mojave Desert
1997,34,
837-846
shrubs
M. RASOUL SHARIFI, ARTHUR C. GIBSON and PHILIP W. RUNDEL
DepartmentofBiologyand theLaboratoryofStructural
and MolecularMedicine,University
of California,Los
Angeles,California90095-1606,USA

Summary
1. Windblowndust,an environmental problemin manydisturbedaridlands,has the
potentialto affectthe physiologicalperformanceof desert shrubs.Physiological
parametersof gas exchangeforthreespecies(Larrea tridentata, Hymenocleasalsola
and Atriplex canescens) weremeasuredat a Mojave Desertsite,at whichbothundis-
turbedand heavilydustedindividualshrubsoccurred.
2. Maximumratesof netphotosynthesis (A) of dustedorganswerereducedto 21%
ofthoseofcontrolplantsin resinousleafletsofLarrea, to 44% in resinousleavesand
photosynthetic stemsof Hymenoclea, and to 58% in non-resinousC4 leaves of Atri-
plex, whichhave vesiculatedtrichomes.Dusted plants of all threespecies showed
reducedmaximumleafconductance(gs), transpiration (E) and instantaneouswater-
use efficiency(A/E). Intrinsicwater-useefficiency(A/gs)was also reduced,exceptin
Atriplex,in whichit remainedunchanged.
3. Temperaturesof dustedleaves and photosynthetic stemswere2-3 'C higherthan
thoseof controlplants,due to greaterabsorptanceof infra-red radiation.Dust also
increasedphotosynthetically
significantly activeradiation(PAR) reflectance.
4. Heavily dusted shrubshad smallerleaf areas and greaterleaf-specific masses,
suggesting thattheshort-term effects
ofreducedphotosynthesis and decreasedwater-
maycause loweredprimaryproductionin desertplantsexposedto dust
use efficiency
duringseasonswhenphotosynthesis is occurring.

Atriplexcanescens,dustpollution,
Key-words: Hymenocleasalsola,Larrea tridentata,
waterrelations.
photosynthesis,
JournalofAppliedEcology(1997) 34, 837-846

and surfacesoilsarelessdry(CaliforniaAirResources
Introduction
Board 1993). Human factors of disturbanceand
Baresoils,whichcharacterize mostdesert ecosystems, increaseddesertification,resultingfromchanges in
area majorsourceofsmallparticulates thatproduce land managementpractices,also have significant
desertdust(Pew&1981).Mediumand
characteristic impactson dustformation(Glantz 1977; Pew&1981;
largesoil grainstypicallymoverelatively shortdis- Grainger1990).
tancesbymodified saltationorshort-termsuspension, The depositionof fineparticulatedust on plant
whereassmallerparticulates (< 20Mm)may enter surfaceshas been reviewedin detailby Chamberlain
long-term suspension andbe transported greaterdis- (1975). Dust grains< 10 /tmin diameterpredominate
tances(Goudie1978;Pye1987).In dryregions subject on plant surfaces,and such deposition frequently
toduststorms, suchas theTharDesertandAmerican resultsin dustclothingshrubsborderingdirtroads or
GreatPlains,aerosolparticleconcentrations may downwindof a barren source area, such as a dry
exceed100/tgm-3(Orgill& Sehmel1976;Goudie lake. For desertshrubsgrowingalongunpavedroads,
1978).Naturalpatterns ofdustmovement oftenvary heavyduston leaves oftenappears to reducethevig-
annuallyand seasonally, as in theMojaveDesertof our of impacted shrubs. Dust deposition in the
California,whereambient levelsoffineaerosolpar- Mojave Desert of Nevada has been shownto cause
ticulates
displaya sharpseasonalcycle,withhighcon- plantdefoliationand shootdeathin Larrea tridentata
centrationsfrequently reaching m-3duringthe
80Mtg (Sesse & Moc. ex DC.) Cov. (Beatley 1965), but the
? 1997 British drysummer monthsbutwithlow concentrations of ecophysiologicalimpacts of dust on desert plant
Ecological Society 10 Mtgm-3during winter,whenplantcoverincreases growthand developmenthave not been quantified.

837

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838 Investigationof theseeffectsmay have implications
Dust effects
on forland managementpracticesin theMojave Desert
desertplants and otherarid and semiaridzones,bothin indicating
how certaintypesof land use, such as urban devel-
opment,road building and militarytraining,may
affectthe natural environmentand in determining
how thenegativeimpactofhumanactivitiesin certain
areas may be minimized.Furthermore,the exam-
ination of naturallyoccurringdust effectscan be
expectedto contributeto our understandingof the
adaptationsthatallow plantsto survivein thedesert
environment.
This field study investigatedthe physiological
effectsof dust accumulationon the photosynthetic
organsofthreecommonMojave Desertshrubs.Creo-
sote bush Larrea tridentata(Zygophyllaceae)is a
widespreadand ecologicallydominantC3 evergreen
shrubwithresinous,bifoliateleaves (Barbour 1969;
Mabry, Hunziker& DiFeo 1977); its leaves show a
highdegreeof toleranceto thelow wateravailability
of its deserthabitat(Oechel,Strain& Odening1972;
Odening, Strain & Oechel 1974; Syvertsen,Cun-
ningham& Feather 1975; Syvertsen& Cunningham
1977; Mooney,BjOrkman& Collatz 1978; Meinzeret
al. 1986,1990b;Sharifiet al. 1988;Lajtha & Whitford
1989; Rundel & Sharifi1993). Burrobrushor chee-
sebushHymenocleasalsola Torrey& A. Gray ex A.
Gray (Asteraceae),especiallycommonalong runnels
and washes,is a C3 shrubwithresinousphotosynthetic
youngstemsand narrow,drought-deciduous leaves.
Because theseleaves are relativelyshort-lived,
photo-
synthesisby stemscontributessubstantiallyto sea-
sonal carbonbalance (Comstock& Ehleringer1988).
FourwingsaltbushAtriplexcanescens(Pursh) Nutt.
(Chenopodiaceae) is a C4 facultativelyevergreen
shrubwithnon-resinousleaves thathave vesiculated
trichomesfor sequesteringsalts fromchlorenchyma
(Osmondetal. 1969;Osmond,Bjorkman& Anderson
1980). Atriplexcanescensis a glycophyte, widelydis-
tributedin aridand semiaridwesternNorthAmerica,
and has received limited ecophysiological study
(Ruess & Wali 1980;Freemanet al. 1993).
It was hypothesizedthatheavydust accumulation
on leaves would have a significantphysiological
impact, as a response to changes in both pho-
tosynthetically activeradiation(PAR) reachingpho-
tosynthetic tissuesand theoverallenergybudgetof a
leaf. This hypothesiswas tested during summer
droughtforthe threestudyspecies,withdustedand
controlplantsgrowingalong an unpavedroad, when
thesensitivityofleafenergybudgetwouldbe greatest.
Materialsand methods
Field studieswere conductedat the United States
? 1997British
Ecological Society, Army Fort Irwin National Training Center/
JournalofApplied Goldstone TrackingStation in the centralMojave
Ecology,34, 837-846 Desert, 50 km north-westof Barstow, California
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(35020'N, 116050'W,elevation 980m), on a gentle
bajada 2km due east of GoldstoneDry Lake. Veg-
etationat thissitewas Larrea-Ambrosiadesertscrub,
which characterizesmost lowland habitats of the
Mojave Desert(Beatley1976;Vasek & Barbour1977).
At thesitewerewidelyspaced shrubsofL. tridentata,
Ambrosiadumosa (A. Gray) Payne, H. salsola and
Atriplex canescens, with scattered individuals of
Acamptopappus sphaerocephalus (A. Gray) A. Gray,
Achnatherum hymenoides (Roemer& Schultes)Bark-
worth,and EphedranevadensisS. Watson. Soil was
alluvialin origin,sandyloam to fineloam and lacking
surfacegravel,and A and B soil horizonscontained
fineparticulateserodedfromthe drylake. This type
of loessial, playa soil is common in the California
deserts.The mean annualprecipitation at thesitewas
170mm.The lowerbajada was crossedby a dirttank
trail used for transporting militaryvehiclesduring
training manoeuvres.A persistent and strongup-slope
windduringtheperiodofspringtrainingmanoeuvres
yieldedan extremely heavydustingon shrubsalong
thedownwind(upper) side of thetanktrailand little
or no detectabledustingalongtheupwind(lower)side
of thesame trail.
Physiologicalparameterswere monitoredin July
1994, during a hot period (midday ambient tem-
peratureoftenexceeding45 C) of summerdrought
and whenno precipitation had been recordedat the
site since the shrubshad been dusted by vehicular
traffic.The studywas conductedduringthe summer
period,because previousobservationsindicatedthat
dustwas onlypresenton leavesduringthatseason. In
winterand spring1993 and 1994 rainfallprevented
dust dispersion.Mature shrubs of Larrea, Hymen-
oclea and Atriplexoccurredon bothsidesof thetank
trail,and foreach speciesfiveto 10 individualswere
selectedfromthedustedand controlareasofthestudy
site.Shrubdensitywas thesame fordustedand con-
trolplots,whichwereseparatedonlyby theunpaved
road and showed no perceptibledifferences in top-
ographyand othersitecharacteristics. The dustedand
control sites were essentiallyhomogeneous except
withregardto dustexposure.
Net photosynthetic rate(A), stomatalconductance
(gs),transpiration (E), instantaneous(A/E) water-use
efficiency(WUE), intrinsicWUE (A/gs)and internal
CO2 concentration (cl) weredetermined underambi-
ent fieldconditionsusing a portablephotosynthesis
system(LI-6200, LI-COR Inc., Lincoln, Nebraska,
USA). Small leafytwigsofeach individual,havingan
averagephotosynthetic surfacearea of8-10 cm2,were
sealed into a 250-mlleaf chamberforgas exchange
measurements. All gas exchangemeasurements were
made from9.00 to 12.00hours,whenphotosynthetic
ratesforall specieswereestablishedto be maximum.
Aftergas exchangemeasurementswere completed,
thosesampleswereharvestedand thesurfacearea was
obtained using a LI-3100 leaf area meter(LI-COR
Inc.). Leaf temperatures inside the cuvettewere not
839 onlymeasureddirectlywiththermocouplesbut also
M.R. Sharifi, calculatedfroma leafenergybalance (Rochetteet al.
A.C. Gibson& 1990). Whereasthe defaultconfiguration of the LI-
P. W. Rundel 6200 does not make thisanalysis,a programwithin
the instrument was modifiedto do this calculation
(LI-COR 1987). Measurementswere also made to
quantify nocturnaltranspiration.Middayshootwater
potentialsweretakenon all sampledindividualsusing
a Scholander-typepressure chamber (n = 10-15).
Reflectanceofleafsurfacesfromdustedand undusted
(control)plantsof Larrea was measuredusinga LI-
1800spectroradiometer (LI-COR Inc.) equippedwith
an integratingsphere to quantify reflectanceat
350-1100nm. Leaf absorptance could not be
measuredwiththisapparatusbecauseleavesweretoo
small.
Quantitativephenology,such as changesin shoot
growthrates and numberof lateralshoots,was fol-
lowedin controland heavilydustedplantsforLarrea
only.Dependingupon canopysize,fiveto 10 terminal
shoots of similarlength(8-11 cm), diameterand age
wererandomlyselectedin each individualin thefour
ordinalcompassdirectionsforphonologicalmeasure-
ments.These shootsweretaggedat theninthor tenth
internodefromtheleadershoottipusingloose-fitting
plasticbirdbands.Total lateralshootlengthand num-
ber of lateralshootsweremeasured.
The amount and distributionof dust on pho-
tosyntheticorgans were determinedfrom samples
adjacentto each shootused forgas exchangebutafter
physiologicalexperimentswere concluded, so that
shootwaterpotentialswerenot affected forgas exch-
ange measurements.For one set of samples,shoot
dustwas removedby repeatedagitationand soaking
in a beaker of distilledwaterfor 24 h, precipitating
dustfor7 days,filtering, oven-drying theprecipitate,
and then weighingdust on a Mettlerbalance. For
these,leafarea was measured.Anothersetof samples
was gentlyair-driedbetweenpapersin a plantpress,to
avoid exposureto dust-removing solvents,and viewed
withan ETEC Autoscan ScanningElectronMicro-
scope at 10 kV aftersputtercoatingwith20 nm of
gold-palladium.For Larrea, 12 adaxial and abaxial
surfacesof companionleafletswere comparedfrom
dustedand controlplants,and forHymenocleadown-
wind and upwind stem and leaf surfaces were
compared.The presenceof rupturedbladdercells on
leaves of Atriplexproducedchargingand thusinter-
feredwith photographicdocumentationof surface
dust. Percentagecover was crudelyestimatedfrom
electron photomicrographs(600 x, 1500 x and
10000 x ) by recordingthepresenceof dustparticles
withinrandomlysampledquadratsof millimetre-rule
graphpaper.
Data wereanalysedstatisticallyby Student'st-test.
Unless indicated otherwise,data are presentedas
? 1997 British
Ecological Society, mean + standarderror(numbersin parenthesesare
JournalofApplied standarderrorsof therelevantmeans;not significant
Ecology,34, 837-846 indicatesP > 0.05).
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Results
On bothsidesofthetanktrail,leafletsofLarrea were
orientatedwithabaxial surfacefacingthe prevailing
up-slopewind(Fig. la, b). Abaxial and adaxial leaflet
surfacesofcontrolplantswerebrightgreenand resin-
ous; unicellulartrichomeswere embeddedin resin,
numerousunobstructedstomata were evident,and
dustparticleswerevirtually absenton adaxial surfaces
(away fromthewind) (Fig. 2a), eitherattachedto the
resinor embeddedwithinit,althoughabaxial surfaces
occasionallyhad 20% of a fieldof view with light
dust. In sharp contrast,heavilydusted leaves were
to a
dull greyon the abaxial surfaces,corresponding
complete,loose coating of dust on the leaf, hiding
trichomes(Fig. 2b), althoughsome greenshowed on
the adaxial surfaces.Many unobstructedstomata
could stillbe observedthroughthe dust cover,and
dust particleswere also embeddedin resin(Fig. 2c);
some wereirregularin shape, and mostwere 1-4 /tm
in diameter(Fig. 2d). Giventhatmaterialsweredried
in a press, therewas no reliableway to determine
percentageinvivodustpluggingofstomata,butdedu-
stedleaves,in whichwaterremovedsurfaceparticles,
had similarstomataldensitiesto dusted specimens.
Washingof heavilydustedleaves produced 16g dust
m-2leaf,or halfthatamounton each surface,exclud-
ingtheamountembeddedin resin.
On controlplants of Hymenoclea,resinousgreen
leaves and stemshad fewor no dustparticleson any
surface,althoughsomewerefoundon downwindsur-
faceswheretheywerelodged along cell margins.On
heavilydusted specimens,downwindsurfaceswere
distinctlygreydue to accumulateddust, especially
along cell margins(Fig. 3a), which also adhered to
theresin;upwindsurfaceswereless affected (Fig. 3b).
Controlplantshad verylittleduston leaves,whereas
on dustedleaves particlescovered > 50% of theleaf
perimeterbut were relativelysparse along the trich-
ome-hiddenmidveinoftheadaxial leafsurface.When
viewedwithSEM, manystomataofHymenocleawere
hidden,occludedby resin,but thiswas consideredto
be an artefactbecause copious resinis excretedfrom
glandulartrichomesduringdryingof thesestemsand
leaves.
a plantthatnormallyhas greyishleaves
In Atriplex,
due to the presenceof vesiculatedtrichomes,dusted
plants were moregreyand releasedsurfacedustwhen
shaken.Dust particleswere attachedto the exposed
surfaceof each bladdercell and occurredamong the
interiorlayersof trichomes.Leaf washingof heavily
dustedshootsproduced40 g dustm-2 leafarea.
Dusted individualsof all threespecies had con-
sistentlysmallermeanleafarea butgreaterleafspecific
mass (Table 1).
Whereasmiddayplantwaterpotentialsin control
plantsof Larrea were0 9 MPa and Hymenocleawere
0 7 MPa higherthandustedplants(P < 0.05), inAtri-
plex waterpotentialin controlplants was 0 8 MPa
lowerthanin dustedindividuals(P < 0 05) (Fig. 4).
840
on
Dust effects
desertplants

N. I

b~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~b

plants
Fig. 1. (a) Larrea tridentata growingon oppositesides of a dustytank trail.Controlplant (no dust), growingdown-
slope and upwindof dust.(b) Heavilydustedplant,growingdownwind,immediately adjacentto dustsource.

Maximumratesof net photosynthesis werefound ameterson waterrelationswas a reductionin tran-

to be sharplyreducedin dusted shoots of all three
species.This reductionwas smallestin Atriplex,with
dustedplantshavingphotosynthetic ratesequivalent
to 58% of those of controlplants (P < 0 05), while
this figurewas 44% (P < 0 01) in Hymenoclea,and
21% (P < 001) in Larrea (Fig. 5a). Maximum leaf
conductancewas similarlyloweredsharply,but pro-
portionallyless thanphotosynthesis(Fig. Sb).
The net effectof changes in gas exchange par-
spirationrangingfrom79% of thatof controlplants
in Hymenocleato 47% in Larrea (P < 0 05) (Fig. 6a).
WUE was loweredsignificantly
Intrinsic (P < 0 05) in
Larrea and Hymenocleabut remainedunchangedin
Atriplex(Fig. 6b). All species showed a significant
(P < 0 05) declinein WUE measuredas the ratio of
A/E (Fig. 6c). The ratioof internalto ambientCO2 in
C4 leaf tissues of Atriplexremainedunchangedat
thealgebraicrelation-
about 0 27 (Fig. 6d), reflecting

Table 1. Morphologicaland physiologicalparametersof control(no dust) and heavilydustedsamplesfromthethreespecies

in thestudy:leafarea perleaf(LA), specificleafmass(SLM), maximumrateofCO2 assimilation(Amax) and maximumstomatal
watervapour conductance(gmax) Values are means obtainedfromfiveto 10 shrubsof each species and treatment group.
N = 100 leavesper speciesand treatment groupforLA and SLM; n = 20-40 measurements per speciesand treatment
group
forAmax and gmax. Numbersin parenthesesare standarderrorsof therelevantmeans.Data wereobtainedin July1994

LA SLM Amax gmax

Species Treatment (mm2) (mgcm-2) ( jmol m-2s-') (mol m-2s-')

Larrea tridentata Control 21 19 50 13 2 0 20

leaves (7) (0-89)
Dusted 17* 24 62 2.3** 0 04**
(6) (1 61)
Hymenocleasalsola Control 7 70 02 10-5 0 17
stems+leaves (3) (28 01)
Dusted 3* 12790 5.8** 0 10*
(1) (50.0)
Atriplexcancescens Control 28 58 19 9 0-14
leaves (5 29)
ig 1997 British Dusted 38 84 12.9* 0-05*
Ecological Society, (5 84)
JournalofApplied
Ecology,34, 837-846 *P < 0 05; **P < 0 01, fordifference
betweentreatments.

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841
M.R. Sharifi,
A.C. Gibson&
P. W. Rundel
14~~~~a

40t,

(a) b

(C) d

Fig.2. Scanningelectronphotomicrographs of leaves of Larrea tridentata.(a) Adaxial surfaceof control leaf, showing
prominent stomataand trichomes embeddedin resincoatingbutessentiallyfreeof particulates.(b) Abaxial surfaceof heavily
dustedleaf;all epidermalfeaturesare hidden,but gaps in dustedsurfaceshow locationsof some stomata.(c) Adaxial surface
of dustedleafin patchwheresurfacedustis mostlyabsentbut particulateshave been embeddedin resin.(d) Abaxial surface
withsome tightly adheredclay particulates.Magnification bars forFig. 2a-b = 25 Mim;
2c, d = 3 Mzm.

iT 1997 British
EcologicalSociety, Fig.3. Scanningelectronphotomicrographs of stemsof dustedHymenocleasalsola plants.(a) Dusted downwindsurfaceon
JournalofApplied whichparticulatesare mostlylodged in depressionsalong cell margins.(b) More lightlydusted upwind surface,having
Ecology,34, 837-846 bar = 25 gm.
particulateswidelydispersed.Magnification

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 -7
842 mately20% whilereflectance of PAR irradiancewas
F Control
Dust effects
on increasedby about 40% (Fig. 8).
desertplants
6o12 _ . Dusted
While phonological measurementsshowed no
0
growthor evena slightdecreasein totalshootlength
c 4 in dusted individuals,in controlplants therewas a
steady increasein total shoot lengthand the total
0
-3 numberof lateralshoots(Fig. 9).
o 2

Discussion

Physiologicalinfluences ofdustaccumulationon pho-

Atriplex Hymenoclea Larrea tosyntheticsurfacesof these desert shrubsparallel
Fig. 4. Midday shoot waterpotential(MPa) of control(no resultsobtainedin previousresearchof dustimpacts
dust)and heavilydustedshrubsofLarrea tridentate, Hymen- on European roadsideplants(Eller 1977; Thompson
oclea salsola and Atriplexcanescens.Barindicates standard et al. 1984). Lowered of dustedsurfacesproduced
g,
error.Data werecollectedin July1994.
changesin leaf energybalance and promotedhigher
leaf temperatures. In leaves of Viburnum tinusL., a
0-1 dust load of lo g m-2leaf reducedPAR absorptance
4 (a) Cotl by20% and caused a 17% decreasein photosynthesis
Ju , 14. (Thompsonet al. 1984). Increasedweightof dustper
12 - Dusted
E unitleafsurfacewas also foundto be inversely related
E to absorptance of PAR, therebysuggestingthat
8 increaseddensityofdustwouldfurther reducetherate
06 of photosynthesis under summerconditionsof high
4~
4 ambientair temperatures. It could be arguedthatan
increasein thedensityof dustwould be beneficialto
a-
a desertplant,if the leaf operatesabove lightsatu-
(b)
0-14- ration,but thiswould not applyin thisstudy,where
ca0*12- the species studiedhave extremelyhigh lightsatu-
E
0*10
rationvalues (Comstock & Ehleringer1988; M. R.
Sharifi,unpublisheddata).
0
E00
Our resultsindicatethat dustedplants had lower
c0-06-
WUE, measured as instantaneousWUE (A/E) or
0-04-
intrinsicWUE (A/gs).A/E,A/gsand CiCa are all com-
ponentsof plant WUE thatreferto thesame physio-
logicalsetpointdetermined bytheinteraction ofpho-
Atriplex Hymenoclea Larrea tosynthetic capacity and the stomatal control of gas
sign (a) Mid-morning averageratesofphotosynthesisfor
control(no dust)andheavily dustedsamplesofLarrea tri- exchange.Meinzer,Goldstein& Grantz(1990a) sug-
dentata,Hymenocleasalsola and Awtriplexcanescens.(b) Mid- gestedthatA/gsshouldprovidemoreconsistentesti-
morning averagevaluesofstomatal conductance forwater
mates of wateruse-efficiency than A/E, because the
vapour.Barindicates standard error.
Data werecollectedin
July 1994. difference between the evaporative demandin theleaf
chamberand thatwhichprevailedbeforesealingthe
leafinsideis immediately reflectedin E, but not in gs.
Several mechanismsmay contributeto the decrease
shipof thiscalculationwithvalues of intrinsic WUE. of WUE in dusted individuals.Flickiger, Oertli &
The ci/ca ratio of dusted shrubs increased in Larrea Fltickiger(1979) foundthatcompleteclosureof sto-
and Hymenocleato levelsofabout 065, and themost mata can be preventedby dust particles.Rawson &
significant (P < 005) changeoccurredin Larrea. Clarke (1988) reportedthatstomataof dustedwheat
Night-time stomatalconductanceand transpiration leaves took severalhours to reach theirmost closed
werelow in bothcontroland dustedplantsof Larrea position, and the currentvapour pressure deficit
(Table 2). This contrastedwithsignificant differences (VPD) had a major effecton theamountand pattern
in daytimestomatalconductanceand transpiration ofnighttranspiration. More fieldand laboratorystud-
(Figs 5b and 6a) betweenthesesetsof plants. ies will be required to elucidate how dust affects
Temperatures ofdustedphotosynthetic organswere physiologicalperformance in thedesertenvironment.
Because greenleaves are almost totallyreflective
of near infra-redradiation(Nobel 1991), it was not
? 1997 British
Ecological Society, surprisingto observe that accumulated leaf dust
JournalofApplied increased absorptance of near infra-red solar
Ecology,34, 837-846 irradianceand therefore caused higherleaf tempera-

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All use subject to JSTOR Terms and Conditions
 _1 0 '140 LIIIcontrol
843 'c 9 (a) (b)
M.R. Sharifi, T 8 120 - Dusted
A.C. Gibson& 0-7
7 100 -
P. W. Rundel E 6 E
[DE 80 -
C: 4 -E 60-
3 - ~~~~~~~~~~~40-
CL 2
(/2 20
C: 1
Atriplex Hymenoclea Larrea Atriplex Hymenoclea Larrea

2.0 -0-8
(c (d)
7 1-5 -0-6-
E
0- 10 0.4-

Uj 0.5 -L 0-2

Atriplex Hymenoclea LarreaAtriplex Hymenoclea Larrea

Fig.6. Effectsof dust on gas exchangeparametersof control (no dust) vs. heavilydusted shoots of Larrea tridentata,
(b) intrinsicWUE (A/g,);(c) instantaneousWUE (A/E); (d) ratio
Hymenocleasalsola and Atriplexcanescens:(a) transpiration;
of internalCO2 concentrationto ambientCO2 concentration (C,/Ca). Bar indicatesstandarderror.Data werecollectedin July
1994.

(15 September)and daytime(16 September)ratesofstomatalwatervapourconductanceand transpiration

Table 2. Night-time
at thestudysiteat GoldstoneTrackingStationduring1994.Numbersin parenthesesare standarderrors
forLarrea tridentata
of therelevantmeans

Daytime Night-time Day/nightratio

9S E 9s E
molm-2s-' mmolm-2s-' molm-2s-I mmolm-2 s s E

Control 0 076 2 49 0 018 0 40 4 31 6 15

(0 009) (0.28) (0.002) (0 05)

Dusted 0 039 2 08 0 0016 0 33 2 35 6 38

(0 008) (0.33) (0-002) (0-03)

50 0-7

LI Control Dusted 06
? so ~Control|
O 054
?: 0 3 usted
a)~345 aQ) 032
E v0.2

00 I I
200 400 600 800 1000 1200

40 Wavelength(nm)
Atriplex Hymenoclea Larrea
Fig.8. Measured ofsolarirradiance
reflectance (wavelengths
ofcontrol(no dust)and
Fig.7.Measuredleaftemperature 350-1100nm)forcontrol dustedleaves
(nodust)andheavily
heavilydusted samples of Larrea tridentata,Hymenoclea inJuly1994.
Data werecollected
ofLarreatridentata.
salsola and Atriplexcanescens.Bar indicatesstandarderror.
in
was42 5 'C at thetimeofdatacollection
Airtemperature
July1994.
1350nm) for dusty leaves compared with control
? 1997British
Ecological Society, to cause a 2-4?C increasein leaf
leaves, sufficient
JournalofApplied tures.Experimentalstudiesin Switzerlandshowed a temperature (Eller & Brtinner1975; Eller 1977). This
Ecology,34, 837-846 doublingof absorptancein the near infra-red(700- matchesthe2 'C increaseobservedinLarrea and Atri-

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All use subject to JSTOR Terms and Conditions
844 (a) waterstress(Ricks & Williams1974;Eller& BrUnner
E 0 Dusted 0 Control
Dust effects
on 0
60 1975). While our resultsfor nocturnalconductance
desertplants do not supportthismechanismin Larrea, thereare
c 50 numerousstudies(Ricks & Williams 1974; Eller &
?40
Brtinner1975; Eveling & Bataille 1984) that have
reportedthatfinedustparticlesoccludestomata,pre-
30 sumablyby loweringleaf conductanceto watervap-
0
our duringthe daytimeand increasingwaterloss at
20
night.Incompleteclosureof stomatawould cause a
(b)
0
o 25 decreasein WUE, make impactedplants more sus-
ceptibleto drought,and exposeinteriorsof organsto
20 increasedoxidantair pollutants,whichin turnmay
have directand indirecteffectson plantperformance.
The potentialfor small particulatesto wedge open
E 1 50
0
stomatalpores certainlyexistsfordesertplants,but
z directobservationsdid notconfirm thatthisoccurred,
5 because closurewas complete
virtually at night,and
July August September thus particulateswedged into stomatal pores are
unlikelyto have any significanteffecton plantwater
loss.
Fig.9. Phenologicaldataforindividuals
ofLarreatridentata The minimum midday plant water potentials
sampledon 7 July,3 Augustand 16 September 1994at
reportedhere are comparable to values previously
thestudysiteat Goldstone TrackingStation.Barindicates
standard error. reported.Midday summerplantwaterpotentialsof-
6 5 MPa werefoundforLarrea growingin theSono-
ran Desert (Nilsen, Sharifi& Rundel 1984), while
plex and 3 0C increase in photosynthetic stems of values of -5 8 MPa (Bamberget al. 1975) and -6 1
Hymenoclea. highAt ambient summer temperatures MPa (M. R. Sharifi,A. C. Gibson & P. W. Rundel,
of 40-45 0C in thecentralMojave Desert,theserela- unpublisheddata) have been reportedforLarrea in
tivelysmallabsolutechangesin leaftemperature may theMojave Desert.Whilehigherwaterpotentialswere
significantly reduceratesof net photosynthesis. Lar- notedin controlplantsthanin dustedplantsofLarrea
rea has a summeroptimumtemperature forphoto- and Hymenoclea,theoppositeeffect was notedin the
synthesisnear 40'C (Mooney et al. 1978). Any C4 plantAtriplex.Pre-dawnwaterpotential,an indi-
decreasein PAR wouldbe expectedto yielda lowered cator of theequilibriumbetweensoil waterpotential
rate of net photosynthesisof dusted organs. In and plants,was notmeasuredat our site.However,in
addition,leaftemperatures approachingor exceeding a companionstudyof Larrea alone at the same site
45 'C havethepotentialto cause significant heatstress and duringthe same monthand year,no significant
and permanenttissuedamage. differenceswere found betweendusted and control
Anotherpotentialphysicaleffectof dust on pho- individualswithregardto pre-dawnwaterpotentials
tosynthetic surfacescould be a change in boundary (-4 7 and -5 0 MPa, for dusted and controlplants,
layerconditionsdue to increasedsurfaceroughness. respectively) (M. R. Sharifi,A. C. Gibson & P. W.
A thickcoating of dust particleson a leaf surface Rundel,unpublisheddata).
theoreticallywould produce a small decrease in The phonologicaldevelopmentof Larrea control
boundarylayerconductanceacross the leaf/airtran- plants showed a steadyincreasein the total shoot
sitionthatwould lowertranspiration rate,thuslead- length and the total number of lateral shoots.
ing to lower evaporativecooling and consequently However, dusted individualsshowed no growthor
increasedleaftemperatures and reducedgrowth(Dar- even a slightdecrease(due to branchsenescence)in
ley 1966; Eveling 1969; Borka 1980). At the same total shoot length.In most C3 shrubsin the Mojave
time,irregularaccumulationof dust would produce Desert,peak growthactivity continuesuntilmid-sum-
changesinturbulence forairflowovertheplantorgan. mer,whileevenC4 plantssuchas Atriplexreachpeak
Probablymoreimportant, heavyduston a leafcould biomassproductionat midsummer (Ackerman,Rom-
also cover a significant percentageof the stomatal ney & Kinnear 1980). However, peak growthand
pores,therebyloweringleafconductanceand causing phenologicalactivitymay shiftby weeks, or even
elevatedleaftemperatures. The methodsemployedin months,towards springor summer,dependingon
thecurrentstudywerenot capable of determining to temperature (Ackermanetal. 1980).
and precipitation
whatextentstomatawereblockedby dust. Hymenocleashoweda similarphonologicalpatternto
Wheredustparticlesare veryfine,individualgrains Larrea. Atriplex,on the other hand, had higher
? 1997British
Ecological Society, have been reportedto occlude stomata,presumably growthratesthanthetwo C3 plantstowardstheend
JournalofApplied lowering leaf conductance during daylight and of July.
Ecology,34, 837-846 increasingwaterloss at nightor underconditionsof The physiological impactsofduston Mojave Desert

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All use subject to JSTOR Terms and Conditions
845 shrubssuggestthat significant short-term effectsof
M.R. Sharifi, reducedphotosynthesis and decreasedWUE maywell
A.C. Gibson& have long-term effectson netprimaryproductionand
P. W. Rundel shrubvigour,as quantifiedherein smallerleaf areas
and greaterleaf specificweight,whenexposedto fine
particulatesalong unpaved roads. Whereas these
resultsare based on studiesat a singlesite,and must
thereforebe extrapolatedwith some caution, we
believe that our conclusionsmay apply broadly to
theexpectedimpactsof duston plantproductivity in
manysystems. Futurestudiesinvestigating dusteffects
at multiplesites should providefurthersupportfor
thesefindings.
Acknowledgements
We thank StephenAhmann,Mickey Quillman and
RichardAguayo fortheirassistancein some phases
of thisstudy.This researchwas fundedthroughcon-
tract65-9104-4-23 withtheUnitedStatesDepartment
of AgricultureSoil ConservationService.
References
Ackerman,T.L., Romney,E.M. & Kinnear,J.E.(1980) Phe-
nologyofdesertshrubsin southernNye County,Nevada.
GreatBasin NaturalistMemoirs,4, 4-23.
Bamberg,S.A., Kleinkopf,G.E., Wallace,A. & Vollmer,A.
(1975) Comparativephotosynthetic productionofMojave
Desertshrubs.Ecology,56, 732-736.
Barbour, M.G. (1969) Age and space distributionof the
desertshrubLarrea divaricata.Ecology,50, 679-685.
Beatley,J.C. (1965) Effectsof radioactiveand non-radio-
activedustupon Larrea divaricataCav., Nevada Test Site.
HealthPhysics,11, 1621-1625.
Beatley,J.C. (1976) VascularPlantsof theNevada Test Site
and Central-Southern Nevada: Ecologic and Geographic
Distributions.Energy Research and DevelopmentTID-
26881. TechnicalInformationCenter,Officeof Technical
Information, Springfield,Virginia.
Borka, G. (1980) The effectof cementdust pollutionon
growthand metabolismof Helianthusannuus.Environ-
mentalPollution,22, 75-79.
CaliforniaAir Resources Board (1993) Blue Sky Report:
1992 AnnualStatistics.CaliforniaAir ResourcesBoard,
Sacramento.
Chamberlain,A.C. (1975) The movementofparticlesinplant
communities.Vegetationand the Atmosphere(ed. J. L.
Montieth),Vol. 1,pp. 155-203.AcademicPress,London.
Comstock,J.P. & Ehleringer, J.R. (1988) Contrastingpho-
tosynthetic behaviorin leaves and twigsof Hymenoclea
salsola, a green-twigged warm desert shrub. American
JournalofBotany,75, 1360-1370.
Darley, E.F. (1966) Studies on the effectsof cement-kiln
dust on vegetation.Journalof theAir PollutionControl
Association,16, 145-150.
Eller,B.M. (1977) Road dust inducedincreaseof leaf tem-
perature.Environmental Pollution,13, 99-107.
Eller, B.M. & BrUnner,U. (1975) Der Einflussvon Stras-
senstaub auf die Strahlungabsorptiondurch Blitter.
ArchivesderMeteorolgie,Geophysik, undBioklimatologie,
? 1997British SeriesB, 23, 137-146.
Ecological Society, Eveling,D.W. (1969) Effectof sprayingplants with sus-
JournalofApplied pensionsof inertdust.AnnalsofAppliedBotany,64, 139-
Ecology,34, 837-846 151.
This content downloaded from 192.231.202.205 on Fri, 17 Apr 2015 03:03:02 UTC
All use subject to JSTOR Terms and Conditions
Eveling,D.W. & Bataille,A. (1984) The effect ofdepositsof
small particleson the resistanceof leaves and petals to
waterloss. Environmental Pollution,36, 229-238.
Flilckiger,W., Oertli,J.J.& FlUckiger,H. (1979) Relation-
ship betweenstomataldiffusiveresistancesand various
applied particle sizes on leaf surfaces.Zeitschrift fur
Pflanzenphysiologie, 91, 173-175.
Freeman,D.C., McArthur,E.D., Sanderson,S.C. & Tie-
demann,A.R. (1993) The influence oftopographyon male
and femalefitnesscomponentsofAtriplexcanescens.Oec-
ologia,93, 538-547.
Glantz,M.H. (1977) Desertification. WestviewPress,Boul-
der,Colorado.
Goudie,A.S. (1978) Dust stormsand theirgeomorphological
implications.JournalofAridEnvironments, 1, 291-310.
Grainger,A. (1990) The ThreateningDesert. Controlling
Desertification. EarthscanPublications,London.
Lajtha, K. & Whitford, W.G. (1989) The effect ofwaterand
nitrogenamendmentson photosynthesis, leaf demogra-
phy, and resource-useefficiency in Larrea tridentata, a
desertevergreen shrub.Oecologia,80, 341-348.
LI-COR (1987) LI-6200 TechnicalReference.LI-COR Inc.,
Lincoln,Nebraska.
Mabry,T.J.,Hunziker,J.H. & DiFeo, D.R. (1977) Creosote
Bush: Biology and Chemistryof Larrea in New World
Deserts.US/IBP SynthesisSeries6. Dowden, Hutchinson
and Ross Inc., Stroudsburg, Pennsylvania.
Meinzer,F.C., Goldstein,G. & Grantz,D.A. (1990a) Car-
bon discrimination in coffeegenotypesgrownunderlim-
itedwatersupply.PlantPhysiology, 92, 130-135.
Meinzer,F.C., Rundel,P.W., Sharifi,M.R. & Nilsen,E.T.
(1986) Turgorand osmoticrelationsof the desertshrub
Larrea tridentata. Plant,Cell andEnvironment, 9, 467-475.
Meinzer, F.C., Wisdom, C.S., Gonzalez-Coloma, A.,
Rundel,P.W. & Shultz,L.M. (1990b) Effectsof leafresin
on stomatal behavior and gas exchange of Larrea tri-
dentata(DC.) Cov. FunctionalEcology,4, 579-584.
Mooney, H.A., Bjorkman,0. & Collatz, G.J. (1978) Pho-
tosynthetic acclimationto temperature in thedesertshrub,
Larrea divaricata.I. Carbon dioxide exchange charac-
teristicsof intactleaves.PlantPhysiology, 61, 406-410.
Nilsen, E.T., Sharifi,M.R. & Rundel, P.W. (1984) Com-
parativewaterrelationsof phreatophytes in the Sonoran
Desertof California.Ecology,65, 767-778.
Nobel, P.S. (1991) Physiochemical and Environmental Plant
Physiology. AcademicPress,San Diego.
Odening,W.R., Strain,B.R. & Oechel, W.C. (1974) The
effectof decreasingwaterpotentialon net CO2 exchange
ofintactdesertshrubs.Ecology,55, 1086-1095.
Oechel,W.C., Strain,B.R. & Odening,W.R. (1972) Photo-
synthesisratesof a desertshrub,Larrea divaricataCav.,
underfieldconditions.Photosynthetica, 6, 183-188.
Orgill,M.M. & Sehmel,G.A. (1976) Frequencyand diurnal
variationof duststormsin thecontiguousUnitedStates.
Atmospheric Environment, 10, 813-825.
Osmond, C.B., Bj6rkman,0. & Anderson,D.J. (1980)
PhysiologicalProcessesin Plant Ecology. Towarda Syn-
thesiswithAtriplex.Springer-Verlag, Berlin.
Osmond, C.B., Luttge, U., West, K., Pallaghy, C.K. &
Shacher-Hill,B. (1969) Ion absorptionin Atriplexleaf
tissue.II. Secretionof ions to epidermalbladders.Aus-
tralianJournalofBiologicalScience,22, 797-814.
Pewe, T.L. (1981) Desert dust: origin,characteristics, and
effectson man. GeologicalSocietyofAmericaSpecial Pub-
lication,186, 1-303.
Pye, K. (1987) Aeolian Dust and Dust Deposits.Academic
Press,London.
Rawson, H.M. & Clarke, J.M. (1988) Nocturnal tran-
spirationin wheat.AustralianJournalofPlantPhysiology,
15, 397-406.
Ricks,G.R. & Williams,R.J.H.(1974) Effects ofatmospheric
846 pollutionon deciduouswoodlands.II. Effectofparticulate
matterupon stomatal diffusionresistancein leaves of
Dust effects
on
Quercus petraea (Mattushka) Liebl. Environmental
desertplants Pollution,6, 87-106.
Rochette,P., Pattey,E., Desjardins,R.L. & Dwyer,L.M.
(1990) Adjustmentsof leaf temperaturemeasurements in
LI-COR 6200 assimilationchamberusingenergybalance
calculations.Agricultural
andForestMeteorology, 53, 149-
156.
Ruess,R.W. & Wali,M.K. (1980) Daily fluctuationsinwater
potentialand associated ionic changesin Atriplexcane-
scens.Oecologia,47, 200-203.
Rundel, P.W. & Sharifi,M.R. (1993) Carbon isotope dis-
criminationand resourceavailabilityin the desertshrub
Larrea tridentata.
StableIsotopesandPlantCarbon-Water
Relations (eds J. R. Ehleringer,A. E. Hall & G. D.
Farquhar),pp. 173-185.AcademicPress,San Diego.
Sharifi,M.R., Meinzer,F.C., Nilsen, E.T., Rundel, P.W.,
W.M., Herman,D.J. & Clark,P.C.
Virginia,R.A., Jarrell,
(1988) Effectof manipulationof waterand nitrogensup-
plies on the quantitativephenologyof Larrea tridentata
(creosotebush)in theSonoranDesertofCalifornia.Amer-
icanJournalofBotany,75, 1163-1174.
Syvertsen,J.P. & Cunningham,G.L. (1977) Rate of leaf
productionand senescenceand effectof leaf age on net
gas exchangein creosotebush. Photosynthesis, 11, 161-
166.
Syvertsen, J.P.,Cunningham,G.L. & Feather,T.V. (1975)
Anomalous diurnalpatternsof stemxylemwaterpoten-
Ecology,56, 1423-1428.
tialsin Larrea tridentata.
Thompson,J.R., Mueller,P.W., Fluckiger,W. & Rutter,
A.J. (1984) The effectof dust on photosynthesis and its
significanceforroadsideplants.Environmental Pollution,
34, 171-190.
Vasek, F.C. & Barbour,M.G. (1977) Mojave desertscrub
vegetation.TerrestrialVegetationof California(eds M. G.
Barbour& J.Major), pp. 835-867. Wiley,New York.
Received27 July1995; revisionreceived28 May 1996

? 1997British
Ecological Society,
JournalofApplied
Ecology,34, 837-846

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