Plant Physiological Ecology

565 Pages·1998·18.687 MB·English

by  Hans Lambers Ph.D., F. Stuart Chapin III Ph.D., Thijs L. Pons Ph.D. (auth.)

Plant Physiological Ecology
Springer Science+Business Media, LLC
Hans Lambers F. Stuart Chapin III
Thijs L. Pons
Plant Physiological
Ecology
With 356Illustrations
Springer
Hans Lambers, Ph.D. F.StuartChapin III,Ph.D.
DepartmentofPlant Ecologyand InstituteofArctic Biology
Evolutionary Biology UniversityofAlaska
Utrecht University Fairbanks, AK99775,USA
3584CAUtrecht, The Netherlands
and ThijsL. Pons, Ph.D.
Plant Sciences DepartmentofPlant Ecologyand
Faculty ofAgriculture EvolutionaryBiology
UniversityofWestern Australia Utrecht University
Nedlands, WA6907,Australia 3584CA Utrecht, The Netherlands
Coverart from Lourens Poorter,UtrechtUniversity.
LibraryofCongressCataloging-in-PublicationData
Lambers,H.
Plantphysiologicalecology/HansLarnbers, F.StuartChapin,ThijsL.Pons.
p. em.
Includesbibliographicalreferencesand index.
ISBN978-1-4757-2857-6 ISBN978-1-4757-2855-2(eBook)
DOI 10.1007/978-1-4757-2855-2
1. Plantecophysiology. I. Chapin,F.Stuart(Francis Stuart),
III. II. Pons,ThijsLeendert,1948- . III. Title.
QK717.L35 1998
571.2-dc21 97-33273
Printedon acid-freepaper.
©1998SpringerScience+BusinessMediaNew York
OriginallypublishedbySpringer-VerlagNewYorkInc.in1998.
Softcoverreprintofthehardcover 1stedition 1998
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9 8 7 6 5 4 3 2 1
Foreword
The individual is engaged in a struggle for existence (Darwin).That struggle
may be oftwo kinds:Theacquisition ofthe resourcesneededforestablishment
and growthfromasometimeshostileand meagerenvironmentand thestruggle
with competingneighborsofthesameordifferentspecies.Insomeways, wecan
define physiology and ecology in terms of these two kinds of struggles. Plant
ecology,or plantsociology, iscentered on the relationshipsand interactions of
species withincommunitiesand the way in which populations ofa species are
adapted to a characteristic range of environments.Plant physiology is mostly
concerned with the individual and its struggle with its environment. At the
outsetofthisbook,the authorsgive their definition ofecophysiology,arrivingat
theconclusionthat itisapointofview aboutphysiology.Apointofview that is
informed,perhaps,by knowledge ofthe realworldoutsidethe laboratorywin
dow.Aworldinwhich,shallwesay,thelightintensityismuch greaterthan the
200to 500llmoi photons m-2s-1 used in too many environment chambers, and
one in which a constant 20°C day and night is a great rarity. The standard
conditionsused inthelaboratoryare usuallyregardedastreatments.Ofcourse,
there isnothingwrongwith thisinprinciple;onealwaysneeds abaselinewhen
makingcomparisons.Theidea, however,thatthelaboratorycontrol isthenorm
isfalseand can lead to misunderstanding and poorpredictionsofbehavior.
Theenvironmentfromwhich manyplantsmustacquire resources isundergo
ing change and degradation, largely as a result of human activities and the
relentless increase inpopulation.This has thrownthe spotlightonto the way in
which these changes may feed back on human well-being. Politicians and the
generalpublicasksearchingquestionsofbiologists, agriculturalists,and forest
ers concerning the future ofourfood supplies,building materials,and recrea
tional amenities.Thequestionstakeonthegeneralform, "Can you predicthow
'X' will changewhen environmental variables 'Y' and 'Z' change?" The recent
experienceofexperimentation,doneathigh public expense, onCO enrichment
2
and globalwarming,isasoberingreminderthatnot enoughisknownaboutthe
underlying physiology and biochemistry of plant growth and metabolism to
maketheconfidentpredictionsthat thecustomerswanttohear.Evenatthelevel
of individual plants, there seems to be no clear prediction, beyond that the
response depends on species and other ill-defined circumstances. On the
v
vi Foreword
broader scale, predictions about the response of plant communities are even
harder to make. In the public mind, at least, this is a failure. The only way
forward is to increase our understanding of plant metabolism, of the mecha
nisms of resource capture, and the way in which the captured resources are
allocatedtogrowthorstorageintheplant.Tothisextent,Icanseenodistinction
betweenplantphysiologyandecophysiology.There are largenumbersofmiss
ing pieces ofinformationaboutplantphysiology-period.The approachofthe
new millennium, then, is a good time to recognize the need to study plant
physiologyanew,bringingtobearthe impressivenew tools madeavailableby
gene cloningandrecombinantDNA technology.Thisbookistobewelcomedif
itwillencourageecologiststocometogripswiththeprocesseswhichdetermine
the behavior of "x" and encourage biochemistry and physiology students to
take a morerealisticviewofthe environmentalvariables "Y" and "Z."
The book starts, appropriately, with the capture of carbon from the atmo
sphere. Photosynthesis is obviously the basis of lifeon earth, and some of the
mostbrilliantplantscientistshavemadeittheirlife'swork. Asaresult,weknow
more about the molecularbiophysics and biochemistryof photosynthesis than
we do aboutany other plant process.The influenceofvirtuallyeveryenviron
mentalvariableon the physiologyofphotosynthesisanditsregulationhasbeen
studied. Photosynthesis, however, occurs in an environment over which the
individual plant has little control. In broad terms, a plant must cope with the
rangeoftemperature,rainfall, lightintensity,andCO concentrationtowhichits
2
habitat is subjected. It cannot change these things. It must rely on its flexible
physiologicalresponsetomitigatetheeffectsoftheenvironment.Atalaterstage
in the book, the focus shifts below ground, where the plant has rather more
control over its options for capturing resources. It may alter the environment
around its roots in order to improve the nutrient supply.It may benefit from
microbialassistanceinmobilizingresourcesorenterinto more formal contracts
with soil fungi and nodule-forming bacteria to acquire nutrient resources that
wouldotherwisebe unavailableorbeyonditsreach. Towarditsclose,the book
turns to such interactions between plants and microbes and to the chemical
strategiesthathaveevolvedinplantsthatassist themintheirstruggleswithone
anotherandagainstbrowsingandgrazinganimals.Theauthorsend, then,ona
firmlyecological note,andintroducephenomenathatmostlaboratoryphysiolo
gists haveneverattempted toexplore.These intriguingmattersremindus,asif
reminders were needed, of "how little we know, how much to discover"
(Springerand Leigh).
DAVIDT.CLARKSON
IACR-Long Ashton Research Station
Universityof Bristol
April1997
Acknowledgments
Numerouspeoplehave contributed tothe textand illustrationsin thisbook by
commentingonsections and chapters,providing photographicmaterial,draw
ingnumerousfigures,and soon.Apartfrommanyundergraduatestudentsand
some anonymous reviewers, we wish to thank the following colleagues: Rien
Aerts,OwenAtkin, Fraser Bergersen,HennyBlom,Ad Borstlap,Tjeerd Bouma,
NeilBridson,LieveBultynck,Marion Cambridge,ZoeCardon,MelissaChapin,
Christa Christley, David Clarkson, Brent Clothier, Joseph Craine, David Day,
MargrietDekker,MannyDelhaize,David dePury, IngeDorr,NeilEmery,John
Evans, Tatsuhiro Ezawa, Alastair Fitter, Eric Garnier, James Graham, Brian
Gunning, Ellis Hoffland, Arian [acobs-Brouwer, Tibor Kalapos, Gamini
Keerthisinghe,Ronald Kempers, Marga Knoester,MarjoleinKortbeek-Smithuis,
John Kuo,CatarinaMata, MargarethMcCulley,John Milburn, Frank Millenaar,
Harvey Millar, Frank Minchin, Rana Munns, Oscar Nagel, John Passioura,
Kristel Perreijn, Carol Peterson, Larry Peterson, Corne Pieterse, Hendrik
Poorter, Lourens Poorter, Pieter Poot, Hidde Prins, Peter Ryan, Peter Ryser,
Ingeborg Scheurwater, Heather Sherwin, Sally Smith, Ichiro Terashima, Aart
vanBel,AdrievanderWerf,LizvanVolkenburgh,RensVoesenek,Suzannevon
Caemmerer, Michelle Watt, Maria Weiper, Mark Westoby, Lynne Whitehead,
and Piet Wolswinkel.
HANS LAMBERS
F.STUARTCHAPINIII
THIJS1. PONS
vii
Units and Conversions
Concentration, amount
Concentrationequalscontentperunitvolume, mass or area
Molarity = mol-l" solution
Molality = molarity X activity
lmol-m'" = lmmol-kg" = 1mM
1umol CO ·m-3 = 0.00243Pa = 2.43kPa (at293K and 101.3kPa atmospheric
2
pressure
1dalton = 1.6605.10-24g
Energy
1] = 1N'm = Lkg-rrr-s" = 1W·s
1W-h = 3.6kw-s= 3.6kJ
1MJ= 0.278kWh
leal = 4.1868J
1kcal = 1.163W·h
Electric conductance
15 = Lohm"
Gas exchange, conductance
1mol-rn-2'S-1 = 0.0224·(T/273H101.3/P) m-s" (T= absolute temperature;
P = atmospheric pressurein kPa)
t mol-mf-s" = 0.024m·s-J (at293Kand 101.3kPa)
1mm-s" = 41.7mmol·m-2·s-1(at293Kand 101.3kPa)
Pressure
1MPa = 106 Pa = 106N·m-2 = 106[-m" (= 10bar,adiscarded metricunit)
Radiation (McCree 1991)
1W·m-2 = 1]-m-2·s-1
1mol photons = 1·8.105J(at A650nm) to 2.7.105J(at A450nm)
1W·m-2(PAR) = 4.6umol (photons) m-2s-1 (daylight,sunny)
ix
x Units and Conversions
1W·m-2(PAR)= 4.2umol (photons) m-2s-1(daylight,diffuse)
1W·m-2(PAR)= 4.6/4.6/5.0umol (photons) m-2s-1(metal halidelamp/
whitefluorescent tube/incandescentlamp)
Water potential
'P.(MPa) = -RIc.(c. = concentrationin molm")
v'P:air(MPa) = -RI/Vwo·lnRH
= partialmolarvolumeofwater
RH (relative humidityofthe air, %) = p/Po·lOO
p = watervaporpressure
Po = saturatedwatervaporpressure
1M = 103mol·m-3 - -2.4MPa (at20°C)
Reference
McCree,KJ.(1981)Photosyntheticallyactiveradiation.In:Encyclopediaofplantphysiol
r.s,
ogy, N.S.,Vol12A,0.1.Lange, Nobel,C.B.Osmond,& H.Ziegler(eds).Springer
Verlag,Berlin,pp.41-55.
Abbreviations
a radius ofa root (a.) or root plusroot hairs (a.)
A rate ofCO assimilation;also total root surface
2
AI foliage area
A light-saturated rateof net CO assimilationat ambient P.
max 2
As sapwood area
ABA abscisic acid
ADP adenosinediphosphate
AM arbuscularmycorrhiza
AMP adenosine monophosphate
APAR absorbed photosyntheticallyactive radiation
ATP adenosine triphosphate
b individual plantbiomass;buffer powerofthe soil
B stand biomass
Cs concentrationof the solute
C nutrientconcentrationin solution;also convectiveheat transfer
C photosyntheticpathwayutilizing3-carbonintermediate
3
C photosyntheticpathway utilizing 4-carbonintermediate
4
Cli initialnutrientconcentration
Cmin solutionconcentrationat which uptake iszero
CN carbon:nitrogen ratio
CAM CrassulaceanAcid Metabolism
CC carbonconcentration
CE carbohydrateequivalent
chl chlorophyll
CPF carbondioxideproductionvalue
d plantdensity; also, leaf dimension
D diffusivityofsoil water
De diffusion coefficient ofion in soil
DHAP dihydroxyacetone phosphate
DM dry mass
DNA deoxyribonucleicacid
e watervaporpressureintheleaf(e ore inSect.2.5ofthechapteron
j; 1
the plant's energybalance) or atmosphere (e.);also emissivityofa
surface
E leaf transpiration rate
f tortuosity
xi