GPP in Tropical Ecosystems (Monteith, 1972)

Solar Radiation and Productivity in Tropical Ecosystems
Author(s): J. L. Monteith
Source: Journal of Applied Ecology, Vol. 9, No. 3 (Dec., 1972), pp. 747-766
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/2401901
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SOLAR RADIATION AND PRODUCTIVITY IN TROPICAL
ECOSYSTEMS*
BY J. L. MONTEITH
Department
ofPhysiology
and EnvironmentalStudies,School of Agriculture,
University
of Nottingham
In thermodynamic terms,ecosystemsare machinessuppliedwithenergyfroman ex-

ternalsource,usuallythesun.Whentheinputof energyto an ecosystemis exactlyequal
to itstotaloutputof energy,thestateof equilibriumwhichexistsis a specialcase of the
FirstLaw of Thermodynamics. The Second Law is relevanttoo. It impliesthatin every
spontaneousprocess,physicalor chemical,the productionof 'useful' energy,which
could be harnessedin a formsuch as mechanicalwork,must be accompaniedby a
simultaneous'waste' ofheat.No biologicalsystemcan breakor evade thislaw. The heat
producedby a respiringcell is an inescapablecomponentof cellularmetabolism,the
cost whichNaturehas to pay forcreatingbiologicalorderout of physicalchaos in the
environment of plantsand animals.
Dividingthe usefulenergyof a thermodynamic processby the totalenergyinvolved
givesa figurefortheefficiency oftheprocess,and thisprocedurehas beenwidelyused to
analysetheflowofenergyin ecosystems. For example,theefficiency withwhicha standof
plantsproducesdrymatterby photosynthesis can be definedas the ratio of chemical
energystoredin theassimilatesto radiantenergyabsorbedby foliageduringtheperiod
of assimilation.The choice of absorbed energyas a base for calculatingefficiency is
convenientbut arbitrary.To derivean efficiency dependingon the environment of a
particularsiteas well as oil the natureof the vegetation,drymatterproductioncan be
relatedto thereceiptof solar energyat thetop of theearth'satmosphere.This exercise
was attemptedby ProfessorWilliamThomson,laterLord Kelvin,in 1852. 'The author
estimatesthemechanicalvalue of thesolar heat which,werenone of it absorbedby the
atmosphere, wouldfallannuallyon each squarefootofland,at 530000 000 footpounds;
and infersthatprobablya good deal more,1/1000of thesolar heat,whichactuallyfalls
on growingplants,is convertedinto mechanicaleffect.'
Outsidetheearth'satmosphere, a surfacekeptat rightanglesto thesun's raysreceives
energyat a mean rateof 1360 W m-2 or 1f36kJm-2 s-1, a figureknownas the solar
constant.As theenergystoredby plantsis about 17 kJpergramof drymatter,thesolar
constantis equivalentto theproductionof drymatterat a rateof about 1 g m-2 every
12 s, 7 kg m-2 per day, or 2 6 t m-2 year-'. The annual yield of agriculturalcrops
rangesfroma maximumof 30-60 t ha-' in fieldexperiments to less than I t ha-' in
some formsof subsistencefarming. When theseratesare expressedas a fractionof the
integratedsolar constant,the efficiencies of agriculturalsystemslie between0-2 and
0 004%, a rangeincludingKelvin'sestimateof 0-1%.
Conventionalestimatesof efficiency in termsoftheamountof solar radiationincident
at theearth'ssurfaceprovideecologistsand agronomistswitha methodforcomparing
plantproductivity underdifferent systemsof land use and managementand in different
* Openingpaperreadat IBP/UNESCO Meetingon Productivity
of TropicalEcosystems,
Makerere
University,
Uganda,September1970.
F
748 Productivity
in tropicalecosystems
climates(Holliday 1966; Penman 1968). Classifying
ecosystemson the basis of energy
conversionis becominga popular formof taxonomybut it contributeslittleto our
understanding of how plantsrespondto theirenvironment. It is necessaryto go a stage
further,relatingefficiencesof dry matterproductionto the physicaland biological
factorsthatdeterminegrowthratessuchas thefractionof radiationintercepted by a leaf
canopy,the irradianceof individualleaves,the diffusion
resistanceof stomataand the
behaviourof thephotochemicalsystem.This paper attemptsto developthisapproach,
using examplestaken fromtropicalecosystemsalthoughthe principlesinvolvedare
generalenoughto be appliedto any standof plantsgrowingin any climate.
COMPONENTS OF EFFICIENCY
The efficiency
e withwhichcrops or naturalcommunities producedrymatteris defined
hereas thenetamountofsolarenergystoredbyphotosynthesis in anyperiod,dividedby
thesolarconstantintegrated overthesameperiod.Thisdefinition allowse to be expressed
as theproductof sevenfactorsdistinguished by appropriatesubscripts:6g determined by
thegeometryof theearthwithrespectto thesun; 6a determined by thetransparency of
theearth'satmosphere;esdetermined bythespectralcompositionof solar radiationand
bytheopticalpropertiesoffoliage;6q relatedto thenumberoflightquantaneededin the
photosyntheticprocess;sl fractionof radiationinterceptedbya canopy;Ed determined by
thefiniterateat whichcarbondioxidemoleculescan diffusefromtheatmosphereto the
surfacesof photosynthetic unitsin greencells; 6r thefractionof assimilatenot used for
respiration.
Then theefficiencyforthewholesystemcan be writtenas
8 = 8 (g, r) = 8gea8s8qeied8r
and theefficiency
of anypartof thesystemcan be determined theappro-
by multiplying
priategroupof consecutivefactors,e.g.
e (s, d) = 6s6q6i6d
Thegeometrical factor6g
Knowing solarconstant,it is possibleto calculatetherateat whichradiantenergy
the
will be receivedoutsidethe atmosphereon a plane parallelto the earth'ssurface.The
factore9is definedas theratioof thisfluxto thesolarconstantintegrated
overthesame
periodand it dependsonlyon latitudeand on season. Fig. 1, derivedfromthe Smith-
sonianMeteorologicalTables,showsthat6g is relatively constantthroughtheyearin the
tropicshavinga biannualoscillationof +001 at theequatorand an annual oscillation
of +0-06 at latitude200. The much largerseasonal change in temperateregions(e.g.
0-07-0-36at latitude500) is a majorfactordeterminingthepatternof worldclimateand
the global distributionof plant and animal species. The annual average value of 6g
decreasesfromabout 0 3 in thetropicsto 0-2 in temperatelatitudes.
Atmospheric transmission
Solar radiationpenetratingtheearth'satmosphereis absorbedand scatteredbygases,
by cloud, and by aerosols in the formof soil and salt particles,smoke,insectsand
spores.Knowingtheamountofwatervapourand ozone in theatmosphere, it is possible
to calculatehow muchradiantenergywouldbe receivedat thesurfacein theabsenceof
J. L. MONTEITH 749
0-3 ~~~~~~~~~~~0
I0
-
20
1z, 0-2
50
J M M J S N
ofgeometrical
FIG. 1. Annualvariation factoreg at threelatitudes
(N or S).
cloud or of particulatesand to expressthisfluxas a fractionof theextraterrestrial

radia-
tionon a horizontalsurface(i.e. 6g timesthe solar constant).
For threetropicalstations,Samaru in Nigeria (11 N 80E), Kinshasa in the Congo
(40S 15'E) and Mugugain Kenya(1'S, 370E)thewatervapourcontentof theatmosphere
was derivedby interpolation frommaps publishedby Bannon & Steele (1960) and the
ozone contentwas assumedto be constantat 3 mm(Robinson 1966). A computerpro-
gramdevelopedby Dr M. H. Unsworthwas used to calculatethetotalfluxof radiation
at the earth'ssurfaceon the 15thday of each monthand thesefluxesare plottedas a
fractionof theextraterrestrialradiationin Figs. 2, 3 and 4. The mean transmissivity of
a clean cloudlessatmosphereis relatively moreconstantthroughtheyearin thetropics
thanin temperatelatitudesbecause thereis littleseasonal changein themean depthof
atmospheretraversed bythesolarbeam.At all threestationsthetransmissivity is close to
Air
"_, > > Aerosol
0-4 E
J M M J S N
I.0? (
Air Air
0-8 ... . ..
' ------. . _. ..--.*---* - _____,-
Aerosol -- ~~ Aerosol-X
Liilll
0-6 - lu
Cloud
04S~f
J M M
11~
J S N J M M J S N
FIGS. 2-4. Annualvariationof atmospheric transmissionfactor6a ( ), meanmonthly
maximaas fraction-of extraterrestrial
flux(---), and calculatedtransmission
factorfor
cloudlessdust-free atmosphere(- .
FIG. 2. Sumaru,Nigeria.
FIG. 3 Kinshasa,Congo.
FIG. 4. Muguga,Kenya.
750 in tropicalecosystems
Productivity
smallerat Kinshasawheretheprecipitablewaterreachesa maximumof
0-8.It is slightly
about 4 cm in Octoberand slightly largerat Mugugawheretheamountof watervapour
absorbingradiationis restricted by thealtitudeof thesite-2000 m.
The averagetransmissivity of the real atmospheresa was determined by dividingthe
monthlymeaninsolationmeasuredat each stationby thecorresponding extraterrestrial
radiation.Figs. 2-4 showthats followsa different patternat each sitedetermined bythe
seasonal distribution of cloud and aerosol. The annual averageof Sa rangesfromabout
0-48 at Kinshasa wherecloud is prevalentthroughthe year to about 0 58 at Samaru
wherethewintermonthsare almostcloudless.
The differences betweenthe amountsof radiationtransmitted by a cloudless,dust-
freeatmosphereand by a real atmosphererepresent theenergyabsorbedand scattered
back intospace by cloud dropletsand aerosol. The largefractionof radiationscattered
forwardsbyparticulates and byisolatedcumulusis includedin thetotalfluxmeasuredat
the surface.The dashed line in each figurerepresentsa tentativeeffortto distinguish
betweentheeffects of aerosoland cloud. It is themaximumradiationreceivedperday in
each monthof theyearaveragedover5 years.In any givenmonth,themaximumradia-
tionwillusuallybe recordedon a day withlittleor no cloud but thiswillnotnecessarily
be a day when the attenuationby aerosol is representative of the whole month.It is
reassuringthatat Muguga (Fig. 2) and at Kinshasa (Fig. 3), theminimumloss of radia-
tionassignedto aerosolinterception occursduringthewettestmonthsof theyearwhen
the atmosphereis likelyto be cleanestand the maximumloss occursduringthe inter-
veningdrymonths(e.g. May to September).The averageannual loss of radiationat-
tributableto aerosol is verysmall at Muguga, about 0.05 of the extraterrestrial flux,
consistentwiththealtitudeof thestation.At Kinshasa,the averageis about 0.1 of the
extraterrestrialflux,comparablewiththe loss at smoke-free sitesin Britain(Monteith
1966).
At Samaru,the partitionbetweencloud and aerosol shownin Fig. 4 is inconsistent
withthelocal climate.DuringthewintermonthsofNovemberto February,thestationis
almostfreefromcloud but is exposed to dust and sand sweptsouthby the Harmattan
wind.The loss of about 0l15 of theextraterrestrial fluxduringthesemonthsshould be
to
ascribedmainlyto aerosol ratherthan cloud.
Wendland & Bryson(1970) drew attentionto the prevalenceof smoke and soil
particlescarriedup fromthe surfaceduringdry seasons in the tropicsand Figs. 2-4
indicatethataerosoland cloud maybe equallyimportant in determining theseasonaland
of
geographicaldistribution radiation at the surface.
Spectralfactores
The processof photosynthesis in greenleaves needs radiantenergyin the waveband
from0 4 to 0 7 Itm,oftenreferred activeradiation',PAR, or
to as 'photosynthetically
simply 'light'.At theearth's surface,thefractionof radiationreceivedwithinthiswave-
band dependson theextentto whichthe solar spectrumis modifiedby absorptionand
scattering and can be calculatedfora givensolar elevationas a functionof the water
vapour and dustcontentof theatmosphere.Calculationsby Moon (1940) showedthat
the ratio of PAR to total radiationin the directsolar beam was between44 and 45%
whenthesun was morethan300 above thehorizonand a figureof 45% has oftenbeen
usedbybiologiststo calculatethereceiptofPAR fromthefluxoftotalradiationrecorded
witha solarimeter. This estimateignoresthe contribution of diffuseradiationwhichis
scatteredby gas moleculesin theatmosphereand whichcontainsa muchhigherpropor-
J. L. MONTEITH 751
tionofPAR thanthedirectbeam. On cleardays,thediffuse componentis visibleas light
fromtheblue skybutevenwhen the skyis overcast,thesurface of theearthreceivesthis
blue lightby transmissionthroughthe cloud layer.When the solar elevationexceeds
400, theestimatedratioofPAR to totalradiationin thediffuse componentis about 60%.
Combiningthe directand diffuse components in appropriate proportions,the ratio of
PAR to totalradiationis thenclose to 0 50 (Monteith1970).
From a seriesof measurements at Cambridge,England,Szeicz (1970) foundthatthe
annual meanratiowas about 0 49, increasingfroma minimumof 0O48in thespringto a
maximumof 0 51 in thewinter.Therewas a tendencyfortheratioto be smalleron very
cleardays (047) thanon verycloudydays (0-51)probablybecause cloud waterdroplets
absorba smallfractionof solarenergyin theinfra-red spectrum.No long-term measure-
mentsof spectralcompositionhave been reported for the tropics but Fig. 5 shows
measurements made at Samaru,Nigeria in October, 1966, using the Rothamsted band-
pass solarimeterdescribed by Szeicz (1967). An average of 050 is probably appropriate
in thetropicsas well as in temperatelatitudes.
55 0
50 - 0
II I I I
08.00 12.00 16.00
Locol time
FIG. 5. Diurnalvariationof radiationin waveband0 35-07 Apm of totalsolar
as fraction
radiation;Samaru,Nigeria,18 October1966.
Chlorophyllabsorbsverystrongly in theblue and redregionsof thespectrumso that

thelightreflectedand transmittedby leavesis predominantly overthe
green.Integrating
whole spectrumfrom0 4 to 0 7 gum, thefractionof PAR absorbedby leaves is usually
between80 and 90Y/,the precisefiguredependingon factorssuch as the amount of
chlorophyll perunitarea of lamina.An averagefigureof 0 85 willbe adoptedhere.
The fractionof whole-spectrum radiationabsorbedby greenleavesis thereforeabout
0 50 x 0-85 = 0-425and thisquantitywillbe denotedby thesymboles.
Photochemical efficiency6q
When PAR is absorbedby cells containingchloroplasts,the efficiency of photosyn-
thesiscan be definedas theratioofenergystoredin theformation ofcarbohydrates to the
absorbed radiantenergy.Most of the absorbed energyis used in biochemicalcycles
involvingmanyintermediate compoundswhichact as carriersof energy.In veryweak
lightwhenthe rate of photosynthesis is limitedonlyby the supplyof lightquanta and
notbythesupplyofcarbondioxidemoleculesto chloroplasts, about20% oftheabsorbed
energyis storedin thefinalproductsofthephotochemical systemand theremaining 80%,
afterbeingused to formintermediates, is eitherrejectedin theformof heator is used in
the formationof highercompoundssuch as proteinsand fats. More precise,but not
752 Productivity
necessarilymoreaccuratefigurescan be derivedby assumingthattheformationof one
moleculeof carbohydrateneedsone moleculeof CO2 and theenergyof 10 lightquanta
it can be shownthat
(Hill 1973).Takingaccountof thespectralcompositionof sunlight,
theaverageenergycontentof one quantumof PAR is 3-6x 10- 19J.The amountof heat
storedin one molecule(CH2 0) is 7-7x 10-19 J. The maximumefficiency eq of a 10-
quantumprocessin sunlightis therefore
Sq = 10 x (3-6x l0- 19) = 0-215.

Compoundssuch as proteinand fatwhichare formedfromcarbohydrate contribute
onlya smallproportionto theweightof mostplanttissuebutbecausetheyare relatively
richin energyit is necessaryto take theminto account whenthe efficiency of photo-
synthesisis estimatedfromthe amountof drymatterproducedby a leaf or a plant.
Because the chemicalcompositionof dry matteris seldom reportedin productivity
studies,Westlake(1963) suggestedthata representativefigureof 16-7kJg- (4 kcal/g)
should be adopted. Althoughthismay be an averageforherbaceousspecies,a larger
valueof about20 kJg- 1is appropriateforwoodyplants(Lieth1968).Valuesin therange
18-20 kJg'- werereportedforgrassesby Hadley & Kieckhefer(1963) and by Wiegert
& Evans (1964).
Adoptinga figureof 16-7kJg to conformwithcalculationsby otherworkers,1 kJof
= 54 mg of drymatter.It follows
solar radiationis equivalentto (0-425x 0 215)/16&7
thatifP is therateofdrymatterproductionin g m-2 h - 1 and I is solarirradianceforthe
wholespectrumin kW m-2 (i.e. kJs- m-2)
P= 54x10-3x3600I= 19-4L
Recallingtheassumptionsthatwereneededto derivethenumericalfactorin thisequa-
tion,P = 20 I is an acceptableapproximation.
Diffusion efficiency
Ed
(i) Singleleaves.The maximumphotosynthetic derivedin thelastsectionwas
efficiency
a theoreticallimitachievedin principleonlywhenthe irradianceand the grossrate of
photosynthesis are verysmall.In thisstate,theaverageconcentration of CO2 molecules
availableto chloroplastsis constantat a valuedetermined bytheconcentration of CO2 in
theair surrounding theleafand bytherespiration rate.Whenthe irradiance is increased
fromnear zero to a largervalue, the intercellular concentration of CO2 must decrease
because of thefiniterateat whichthemoleculesare transported to thechloroplastsby
diffusionfromthe externalair and fromrespiringmitochondria.This decreasein the
availabilityof intercellularCO2 is responsibleforthecharacteristic shape of thephoto-
synthesis-light curveshownschematically in Fig. 6. As the irradiance tendsto zero,the
grossrateofphotosynthesis perunitincident radiationtends to a constant value shownin
the figureas a limitingslope l/band already derivedin termsof carbohydrateas
20 gh-' kW- '. As theirradianceincreases,thephotosynthesis ratefallsfurther andfurther
1
below themaximumrateof /band approachesa limitingvalue shownas 1/a whenthe
intercellular concentration of CO2 approacheszero. In thisstate,usuallyreferred to as
'lightsaturation',therateofphotosynthesis is proportionalto theconcentration of CO2 in
the external atmosphere.Values of 1/adetermined forcrop plants,exposed to air at 300
vpmof C02, rangefromabout 2 g CH2O m-2 h- 1forcotton,tobacco and sugarbeetto
about 8 g m-2 h- 1 forspeciessuchas maize and sugarcane whichforma distinctgroup
J. L. MONTEITH 753
of photosyntheticallyefficient
plants (D'Costa & Milburn 1973). By introducingthe
appropriatenumericalfactor,it can be shownthatthetotalresistanceto thediffusion of
carbondioxidefromthe externalair to thechloroplastsis 14a s cm-' where1/a is in g
CH20 m-2 h-' so the values quoted fora are equivalentto resistancesrangingfrom
about 7 s cm-' forthe least efficient
groupof speciesto about 2 s cm-' forthe most
efficient.
As theminimumstomatalresistancesreportedfora wide rangeof speciesare
about 1-2 s cm-', thesefiguressuggestthatstomatalcontrolof photosynthesis is likely
to be moreimportant in plantssuchas maize and sugarcane thanin lessefficient
species.
Light
0.
C) I Dark
E
_ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ resp. Light
C
~~~~~~~resp.
FIG. 6. Carbondioxideexchangeof isolatedleafas function EA represents

of irradiance:
measurement ofnetuptakewithinitialslopeb-1 and asymptote a 1; OE is darkrespira-
tion;ED is hypotheticallightrespiration.
Two limitshave now been establishedfor rates of photosynthesis by singleleaves:

P 1/b in veryweak lightand P = 1/a in verystronglight.Analysisof laboratory
measurements byGaastra(1963)andothersconfirms thattheseratescan be combinedin a
simpleinterpolationformulawhichallowstherateof photosynthesis to be expressedas a
functionof irradiance(up to 400 W m-2 PAR) and of carbondioxideconcentration (up
to 300 vpm).The formulais
P = (a+b/I)-l (1)
satisfying theconditionsthatP-+I/bas I-+O and thatPF- 1/aas I-+ oc (Monteith1965a).
WhenP and 1/aare expressedin g CH20 m-2 h-' and Iis in kW m-2, b = 0 05.
Eqn 1 has beenusedin a numberofequivalentformsto estimate
(ii) Plantcommunities.
thetotalphotosynthesis of all theleaves assembledin thecanopyof a uniformstandof
vegetation.Some recentmodelsof canopyphotosynthesis includeso manyparameters
thattheycannotbe used withoutthehelpof a computerbuttheyare rigorousenoughto
predictchangesof photosynthesis ratethatmightoccurin responseto smallchangesin
the area or dispositionof the foliage.In the presentcontext,a relatively simplemodel
(Monteith1965b)can be used to derivethediffusion efficiencyof a standby expressing
thepredictedrateof photosynthesis whenthediffusion resistanceof leaves is finiteas a
fractionof the ratewhenthe resistanceis zero.
The firststepin thismodelis to calculatethearea of leavesexposedto directsunlight.
Assumingthat a layer of leaves containingunit leaf area index interceptsa fraction
(1 -s) of theradiationincidenton thelayer,it can be shownthatin a standwitha total
leafarea indexL thearea indexof leavesexposedto directsunlightis
AO = (I -sL)I(1 -s).
754 Productivity
The averageirradianceof theseleavesis (1 -s)I whereI is theirradianceof a horizontal
leaf.Assumingthatall theseleaveshave thesamephotosynthesis lightcurveand thatthe
carbondioxideconcentration is uniformthroughthecanopy,thetotalphotosynthesis by
sunlitleaves is then
PO = AO(a+b/(l-s)I)-'. (2)
A fractionX of the radiationintercepted
by a leaf is assumedto be transmitted and is
available forphotosynthesisby lowerleaves; but radiationtransmitted by two or more
leaves is neglected.Making an arbitraryassumptionabout the distributionof trans-
mittedradiation,it can be shown that the leaf area index of leaves exposed to light
transmitted throughone higherleafis
Al = {1-sL-(1-s)LsL-1}I(1-s)
and thephotosynthesis
of theseleaves is
P1 = A 1(a + b/IT(-s)I.) (3)
The secondstepin themodelis theintegration
of eqns 2 and 3 overthehoursof day-
lighth assumingthattheirradiancechangessinusoidallyand reachesa maximumvalue
of I* at noon, i.e.
I= I* sin(-)
h
wheret is timeaftersunrisein thesame unitsas h. The integralis a transcendental
func-
tion containingthe six parametersI and h specifyingthe radiationclimate,X and s
specifying the relevantoptical and geometricalpropertiesof the canopy,and a and b
describingtheslope of thephotosynthesis lightcurve.
The diffusionefficiency for a canopy Ed can now be definedas the photosynthesis
calculatedfromeqns 2 and 3 whena is assignedan appropriatefinitevaluedividedbythe
photosynthesis whena is zero,i.e.
_S[Ao{(l-s)a+b/I} '+A1{(l-s)a+b/'I}-'] dt (4)

8d =-(1
+Ta) fh (Ilb) dt
In thelimitwhenthe leaf area indexis verylarge so thatthe canopyintercepts all the

lightfallingon it, sL and 5L-1 tend to zero so thatA0 and A1 tend to (1 -s'1. The
denominatorof eqn 4 can be written(1 + T) Tib whereT is the amountof visiblesolar
radiationreceivedin 1day.If T is expressedin MJm- 2, T = 0-07and 1/b= 20 gh- 1kW- 1
the denominatoris (20 Tx 1-07x 103)/3600or 6 T g m-2, representing the grossdaily
productionof carbohydrate whena = 0.
Fig. 7 showsEd plottedas a functionof T forthreesets of values of a and s selected
fromthe literaturebecause theyare relevantto latercalculations.Line I (a = 0O125;
s = 0 55) is appropriateformaize and sugarcane; line II (a = 0-25; s = 0 65) forrice,
wheatand barley;and line III (a = 0-125; s = 0 55) fordicotyledonssuch as tobacco,
cottonand groundnuts. The relationbetweenEd and dailyradiationis muchmoresensi-
tiveto thevalue chosenfora thanto thevalue of s.
The maximumgrossphotosynthesis fora crop withknowncoefficients a and b can
now be estimatedeitherdirectlyfrommeasuredvalues of solar radiationor froman
appropriatefractionof thesolarconstant.In thefirstcase, thedailytotalof visiblesolar
radiationT MJ m-2 is obtainedfroma solarimeter recordand thediffusion efficiencyis
J. L. MONTEITH 755
Cci Cm-2 day-'
200 400 600
0.8 I I I
0-2_
I0 20 30
MJ m-2 day-'
Ed ofcropcanopiesintercepting
efficiency
FIG. 7. Diffusion For crop
radiationcompletely.
typesI, II, III see text.
fromeqn 4 or fromtheappropriate
estimated curvein Fig. 7. The requiredrateof
is then6 Ed T g CH20 m-2 day-1.
photosynthesis
efficiency
Interception Si
Theinterception efficiency Siofa standcanbe defined as theratiooftheactualrateof
grossphotosynthesis to themaximum rateestimated fora standofidentical plantswith
enoughleavesto intercept all theincidentlight.Fieldmeasurements ofphotosynthesis
usingchambers containing wholeplantsshowthatthephotosynthesis of maizeand
cottonis proportional to theintercepted radiation (Hesketh& Baker1967).It has also
beenfoundthattherateat whichsoybeans andmaizeproducedrymatter inthevegeta-
tivephaseofgrowth is proportional tothefraction ofintercepted light(Shibles& Weber
1966;Williams, Loomis& Lepley1965).Thesemeasurements suggestthatSi can be
identifiedwiththefraction of incidentlightintercepted bya stand,a quantity readily
obtainedbydirectmeasurement or byinference fromtheleafareaindex.
Thephotosynthesis rateofa cropcanopycannotbe strictly proportional to thefrac-
tionofintercepted radiation unlessall theleavesareworking at thesamephotochemical
efficiency.Thisimplieseither(i) all theleavesareworking on thelinearportionofthe
photosynthesis/light curve;or(ii) all theleavesareexposedto thesameradiant fluxand
respond tolightinthesameway.Condition (i) is satisfied inweaklight, i.e. on dulldays
andat sunrise andsunseton bright days.Condition (ii) is satisfied whenphotosynthesis
byshadedleavescanbe neglected incomparison withphotosynthesis bysunlitleaves(or
portion ofleaves)withthesamelightresponse curve.Proportionality observed between
photosynthesis andintercepted radiationcan be ascribedto theoperation ofcondition
(i) (butnot(ii)) in cloudyweather and condition (ii) (butnot(i)) in cloudlessweather.
Proportionality willbepreserved overanyperiodduring whichtherelative proportions of
sunnyand cloudyweather are relatively
constant fromdayto dayand weekto week.
A linearrelation between photosynthesis andintercepted radiation is consistent with
themodeldescribed inthelastsection.Thefraction ofradiation transmitted byunitleaf
layeris s+ (1 -s)r so thefraction intercepted bya leafareaindexofL is
1-{s+(1s-)T}L = 1 _L _[LsLl(1 S)T].
Whentheradiation i) b/Iis muchlargerthan(1 -s)a andit

fluxis veryweak(condition
can be shownfromeqns2 and 3 that
756 Productivity
e = I-s'_[Ls'-1(1-S)T1(1 + T)]
a quantityalmostidenticalto thefractionofintercepted radiationwhenX = 007 because
the termsin square bracketswill usuallybe muchsmallerthan 1- sL. Condition(ii) is
consistentwiththe model for leaves transmitting a negligiblefractionof PAR. Then
puttingX = 0, it can be shownthatthefractionof transmitted radiationand 8, are both
equal to (1 - L). WhenX = 0 07, ratesof photosynthesis predictedbythemodelincrease
almost linearlywiththe fractionof interceptedradiationexceptwhens is verylarge
(e.g. 0 80) and a is small(e.g. 0X120).Thus to a good approximation valid formostcrops
and climates,theratioof actualphotosynthesis bya standto themaximumrateachieved
at fulllightinterception can be calculatedas
Si = 1-[s+(I-s)rT]L.
Fig. 8 showsSi plottedas a functionofs and L whenX = 0 07.
1.0-
0.8
0-6-
0-4-
022_ ~~~~I ~ ~I ~ I
2 4 6 8
LAI
FIG. 8. Interception
efficiency
ej as functionof leafarea indexLAI and thegeometrical
factors (besidecurve).
Respiration factors,
In previoussectionsit was tacitlyassumedthatthegrossuptakeof carbondioxideby
a leafcan be measuredas a functionof theirradianceand of theexternalcarbondioxide
concentration.In practice,the exchangeof carbon dioxide usually measuredin the
laboratoryis a netexchange,thedifference betweengrossphotosynthesis and respiration
rates.If respiration proceededat thesame ratein thelightand in thedark,thetruerate
of grossphotosynthesis could be readilydetermined by addingthedarkrespirationrate
to thenetphotosynthesis ratein thelight.This classicalmethodof estimating grossrates
of uptakeis open to criticism becausean important componentof therespiratory system
in leaf cells is coupled to the photosynthesis system.This componentproducescarbon
dioxide at a rate that increaseswith the photosynthesis rate and thereforewith the
irradiance.True rates of gross photosynthesis are verydifficult to estimatebecause
unknownamountsofcarbondioxideand oxygenmaycyclebetweenthesitesof assimila-
tion and respiration withincells.
Fortunately, it is not essentialto distinguishbetweenlightand dark respirationin
productivity studiesprovidedthenetrateof photosynthesis is estimatedby a consistent
conventionshownin Fig. 6. At an arbitrary irradiance,thenetor apparentrateofcarbon
dioxideuptakeis AB. Respirationat thesame irradianceis BD decreasingto OE in the
dark (whetherBD shouldbe regardedas lightplus darkor simplyas lightrespiration is
stillbeingarguedbyphysiologists: thepointis irrelevanthere,howeverimportant it may
J. L. MONTEITH 757
be biochemically).The true gross photosynthesis is AD, but the classical methodof
estimating grossphotosynthesis yieldsthesmallervalue AB + BD = AC. The netrateof
photosynthesis fromwhichthe net rate of drymatterproductioncan be calculatedis
determined eitherbysubtracting thetruerateof respiration in thelightBD fromthetrue
rateof grossphotosynthesis AD or by subtracting thedarkrespiration rateOE fromthe
grossphotosynthesis in thelightestimatedas AC. The grossphotosynthesis referredto in
previoussectionscorrespondsto AC. Note thattheasymptote1/ais measuredfromthe
axis EC and not fromOB.
The respirationfactorsrcan now be definedas theratio(P - R) :P = 1- RIP whereR
is the weightof carbohydrate used forrespirationper day calculatedby multiplying the
mean dark respirationrate (g m-2 h-') by 24; and P is the weightof carbohydrate
producedby photosynthesis overthesame periodcalculatedby addingthedarkrespira-
tion rateto thenet uptakeof photosynthesis hour by hourthroughout theday. In any
plantcommunity, theratioofrespiration to photosynthesis overthesameperioddepends
on manyinternaland externalfactors,notablythegrowthrateof thestand,thefraction
of totaldryweightrepresented by leaves and otherphotosynthetic organs,and thetem-
peratureof respiring tissue.Accurateand relevantestimatesof 8, are rarebecause plant
physiologistshave seldomtriedto measuretherespirationrateof whole plantsand be-
cause crop ecologistsunderestimated the importanceof respirationin determining net
rates of dry matterproduction.The traditionalfigurefor 8, is about 0 20-025 but
Gaastra (1963) quoted a rangefrom0 25 to 0 50 consistentwithgrowth-room measure-
mentson barley(Watson & Hayashi 1965) and Hamil grass(Ludlow & Wilson 1968),
and withfieldmeasurements on lucerne(Thomas & Hill 1949),barley(Monteith1968),
maize and wheat(Lemon 1970). An evenhighervalue of 0 75 was reportedfortropical
rainforest(Kira et al. 1967)and a similarfigurewas derivedby Muller& Nielsen(1965)
fora forestin thehumidregionof theIvoryCoast.
The proportionof assimilateused forrespirationis expectedto be largerin tropical
thanin temperateclimatesbecause the temperature coefficient forrespirationof green
tissueexceedsthecoefficient forphotosynthesis overtherangeof temperatures in which
plantsusuallygrow.From laboratorymeasurements on isolatedsectionsof tissue,the
Q1o of respiration is close to 2 but Chang (1968) analysedThomas and Hill's measure-
mentson lucerneto give
Sr = 0 92-0 012 T,
whereTis in 0C, givinga Q10 of about 1-4at 20? C.
McRee (1970) recentlyshowedthatthe respirationof a stand of whitecloverin the
vegetativestageof developmentwas a linearfunctionboth of thephotosynthesisrateP
and of thestandingdryweightW. At a temperature of 200 C therelationwas
R = 0-25P+0-015 W(g CH2O m-2 day- 1).
Assuminga Q10 of 2 and dividingthroughby P therelationbecomes
Br= 1- {00625+O000375 WIP}2T/1,

and at 250 C, an appropriatedaily mean temperaturein the tropics,Bris 0 65 -0021
WIP. Duringthevegetative stageof growth,photosynthesis
by a standofdicotyledenous
plantsmayreach P = 50 g m 2 day- 1 when =
W 500 g-2 so thatBr= 0 44.
Takingaccountof thewide rangeof values quoted in theliteratureand therelatively
758 in tropicalecosystems
Productivity
rA
o ao
0~~~~~~~~~
.o
0~~~~~~~~~
O 0
o0
0>-ry_
W5
a 2 E _, 8 a)~~~~~~~~~~~~~~~y
E _4 a U,
C'
0 - U, 0
Cd
0~~~~~~~~~~~~~~0
0
40 d n d
~O o o~ o r
LO00
a.) 10
a)0 ~~~~Q"Oa)
Cd'0
# ~ ~ a
NN
U~~~~~~~~~~~~~~~~~~~~U
0 ~~~~~~~~~~~O
* a'CI a'Cj
J. L. MONTEITH 759
highmean temperatures recordedin the tropics,it was decided to adopt an arbitrary
meanvalue of s = 05 and to adjustthisvalueifexperienceshowedthatit was incorrect
fora particularspeciesor fora particularperiodof development.
TESTS OF THE ANALYSIS

The efficiency withwhicha stand of vegetationis expectedto store solar energyby
photosynthesis can now be derivedas a productof the sevenfactorsexaminedin pre-
vious sections.The percentageaccuracyof e (g, r) or of theequivalentrateof drymatter
productionis thesumof thepercentageaccuracyin theindividualfactorssg,Ea. . . , etc.
The firstfactorsg can be calculatedto anyrequireddegreeof accuracyand maythere-
forebe regardedas errorless. Whensolarradiationrecordsare available,Sa can be calcula-
tedfromvaluesofinsolationand from6gwithan accuracylimitedonlybytheperformance
of the recordingsystem.For a properlycalibratedand well-maintained station,the
uncertainty in sa will be about +2%. At siteswhereit is necessaryto estimatesa from
recordsofcloudinessor sunshinehours,theuncertainty in monthlyaveragesof6a willbe
of theorderof + 10%.
The spectralfactorga iS unlikelyto changemuch,eitherwithclimateor withspecies,
and willrarelydepartfromthe normof 0 425 by morethan + 5%. The photochemical
efficiencysq = 0-215was based on a constantspectralcompositionforsunlight and on an
arbitrary choiceof 10 quanta permolecule.Uncertainty in thisfigure
can be setat + 20%
but if eh is over-or under-estimated, therewill usuallybe a complementary errorin
estimatingEd. In verybrightlight,the two errorswill be equal and opposite(because
photosynthesis ratesdependonlyon a and not on b) and in verydim lightthecomple-
mentary errorin Ed willbe zero (becausephotosynthesisratesdependonlyon b and hence
on the figureassumedfor quantumneed). A figureof + 5% will be assumedfor the
averageerrorin she
The factorSiis calculatedas a functionofleafarea indexL whichcan usuallybe deter-
minedto about + 10% withcarefulsampling.The corresponding errorin si willbe + 10%
whenL is small,so thatSi is proportionalto L, decreasingto zero whenL is largeand si
tends to unity.For the whole life of a crop, the average errorin si may be about
?5%.
Uncertainty in s (g, i) is therefore+ 10+ 5 + 5 + 5 or + 17%, and is a combinationof
randomand systematic errorsin theindividualcomponentsof efficiency. The errorsin
Ed and grare moredifficult to establisha priori,buttheerrorin e (g, r) can be determined
empirically by comparingmeasuredratesof drymatterproductionin thefield,Cmwith
estimatedratesCe. Theseestimatesare derivedfroma figurefore (g, i) expectedto have
an errorof + 17%, froman arbitrary figureof sr = 0 50, and froma value of Ed that
dependson theaccuracyand relevanceoflaboratorydeterminations ofa and b. Compari-
son of Cmand Ce wereattemptedforthefewtropicalcropsforwhichdeterminations of
C and of L wereavailable. To includemoredata, the comparisonwas extendedto re-
centlypublishedmeasurements on tropicalcrops grownin Japan for an IBP project
(Table 1).
Whenpreliminary calculationsshowedthatestimatesof crop growthrateweremuch
moresensitiveto differences in a thanin s, itwas decided(i) to workwiththreevaluesofa
spanningthe range derivedfromphotosynthesis curves publishedin the literature;
(ii) to assign one of thesevalues to each crop; and (iii) to associate one appropriate
value of s witheach value of a as follows.
760 Productivity
a s Crops
(m2 h g-')
GroupI 0-125 0 55 Maize,bulrush millet
GroupII 0-25 0-65 Sorghum, rice
GroupIII 0 50 0 55 Soybean,groundnuts
GroupI contains thetropical grasseswhichareknownto haverelatively fastratesof
photosynthesis in brightlight.Sorghum wouldbe expectedto belongto thisgroupbut
wasassigned to GroupII whenitwasfoundthata cropinNigeriaproduced drymatter
at a rate20-25%lessthantheratespredicted fora GroupI crop.GroupII contains
riceas wellas sorghum andwouldprobably be an appropriate
groupforcerealssuchas
wheatandbarley which, inbright assimilate
light, carbondioxidefaster
thandicotyledons.
GroupIII containstwodicotyledons and is an appropriate
groupforcotton,cassava,
beansand manyothertropical plants.
Parameters assumedto be thesameforall six specieswereb (0 05), X (0.07)and 8,
(0 50). Mean valuesofinsolation werecalculatedfromdailysolarradiation exceptat
Katherine where, in theabsenceofrecords, valueof23 MJdaywasassumed
a constant
through theseason.The leafarea indexwas assumedto increaseexponentially with
timeso themeanvalueofleafareaindexL between twoharvests
wascalculated fromthe
standard formula
L = (L2-Lj)/ln(L21L1).
Resultsof thisexerciseare presented in Figs.9-13 whereeachpointrepresents the
relationbetween Ce and Cmfora periodofbetween 2 and4 weeksduring thevegetative
stageof growth. The distribution ofpointsin eachfigure is determined mainlybydif-
ferencesin leafarea rather thanin solarradiation (cf.changesof E withradiation in
Fig.7 and ofSiwithL in Fig.8).
For maizeand millet(Fig.9), Ce and Cmagreeverywellup to 300g m2 week , a
raterarelyexceeded inthefield.Theestimated ratesofsorghum (Fig.10)tendtobe a few
percentlargerthanmeasured ratesbutdifferences in totaldrymatter production be-
tweenthetallindigenous variety andthedwarf
Fara-fara hybrid NK 300canbe accoun-
tedforpurelybydifferences in leafareaindex.Resultsforfourvarieties ofricegrown
at threesitesshow more scatterthan othercrops(Fig. 11) and thereis a tendency for
to
Ce underestimate the rate of dry matterproduction by mature stands. Thereis less
scatterforfourvarieties ofsoybean(Fig. 12) grownat twofertiliser levelsand at four
different
sites.Exceptionally good of
estimates cropgrowth were derivedfortwovarieties
of groundnuts (Fig. 13) eachin twotreatments: sprayedwithDithaneas a protection
againstCercospora leafspot;andunsprayed. Forthiscrop,differences ofleafareaindex
between varietiesandtreatments accountalmostcompletely fordifferences indrymatter
production during vegetative growth.
Thesuccessofthesecomparisons thattheerror
implies inestimatingdrymatter produc-
tionfromconstant valuesof sr and a duringvegetative growth and fromvaluesof s
calculatedfromleafareaindexis comparable withtheerrorin conventional measure-
mentofproduction, about+ 10-15%overa periodof1 or2 weeks.Duringthereproduc-
tivestageofgrowth, Ce wasalwaysgreater thanCm,probably becausethedevelopment
ofa non-photosynthetic storageorganmadeSr largerthan0 5 and becausesenescence
ofleaveswas responsible fora systematicdecreaseof 1/aafternewleavesstoppedap-
pearing.
Ce forothercrops,Fig. 14 showsmaximum
As an aid to determining weeklydry
J. L. MONTEITH 761
300 .
200 -
200~ ~~~
'0
200 0
0 0~~~~~0
I00 200 300 100 200
200 -
0 O/ 100 /A
100 200 100 1OO

Cm
FIGS.9-13. Comparisonof drymatterproduction ratesfrommeasurements
Cmand esti-
matesCe. For cropdetails,see Table 1.
FIG. 9. *, Maize; 0, bulrushmillet.
FIG. 10. Sorghum: 0 Fara-fara; *, NK 300.
FIG. 11. Rice.
FIG. 12. Soybeans.
FIG. 13. Groundnuts:
circles,S 38; triangles,
F 439; solidsymbols,
sprayed;opensymbols,
unsprayed.
0)~ ~ ~ ~~C
cal cm-2day'?
FIG.~//
9E Maze 0-25rshmllt 0
200 400 600
II I /
300 -/ It
0)/
0 *0 20 30
MJm-2day-'
FIG. 14. Estimateddrymatterproduction
forcropcanopiesintercepting
radiationcom-
pletely.
762 Productivity
matterproductionforeach groupwhene1 = 1. The value of Ce is foundby readingoff
a maximumratefromthisfigureand multiplyingby ei determined fromeqn 4 or Fig. 8.
GENERALIZATION AND EXTENSIONS

Primaryproduction
Having establishedand testeda methodof predictingcrop growthrates whichis
consistentwith fieldmeasurements in the tropicsand sub-tropics,differencesin the
efficiency
withwhichdifferent standsuse energycan now be assignedto specificphysical
or biologicalfactors.Table 2 illustrates
thistypeofcomparisonfortheannualproduction
of drymatterby a Class III crop growingin thetropicsand in a temperate climate.
Table 2. Annualproduction
of drymatter
Tropical agriculture Temperate
Intensive Subsistence agriculture
8e 0 30 0 30 0-20
8a 0 50 0 50 0 35
8L 0.20 0 05 0-33
es 0-425 0-425 0-425
eq 0-215 0-215 0-215
Ed 0 30 0.15 0 35
8r 0 50 0 50 0 60
e (i, r) 0-27% 0034%4 0-50%
e (g, r) 0-041% 0O005% 0 038%4
Ce 11 09 10
(kg Mr-2 year 1)
Table 3. Maximumratesof drymatterproduction

Tropical Temperate
(wet season) (summer)
Class I Class III Class III
8e 030 035
8a 0 60 0-40
El 0095 095
es 0-425 0-425
eq 0-215 0 215
Ed 0-62 0-32 0-36
8r 0050 0 50 0 60
e (i, r) 2.Rio 1-4% 1-9%o
e (g, r) 049%4 0-25% 0-27%
Ce 34 18 19
(g m- 2 day 1)
Differences of latitudeand cloudinessare takenintoaccountby Eg and sa respectively.

The interception factoris largestforthetemperateclimatebecauserelatively littleradia-
tion is receivedduringthe wintermonthswhen the leaf area is zero. In the tropics,
however,large amountsof radiationare receivedthroughout the yearbut can be used
onlywhenthetemperature and rainfallregimeallow a canopyto develop.Growthof the
subsistencecrop is assumedto be limitedby a seriousshortageof nutrients and water.
The maineffectof theseshortagesis to restrict leafdevelopmentso ei is set at 005, and
assumingthatphotosynthesis is also checked,Edis set at 020. A respirationfactorof
060 is assumedfora temperate climateto allow forlowermeantemperatures. The lower
J. L. MONTEITH 763
partof thetableshows: (i) theefficiency of thewholesystemE (g, r); (ii) thecorrespond-
ing rateof drymatterproduction;and (iii) thequantityE(i, r), theefficiency withwhich
solar radiationincidenton a crop canopyis convertedto drymatter.The figuresin the
table and othersthatcan be calculatedfordifferent periodsof growthand systemsof
management are consistentwiththeestimatesof efficiency whichPenman(1968) derived
fromFAO worldrecordsof crop production.
Maximumefficiencies of photosynthesis in a givenclimatecan be calculatedon the
assumptionthatei = 0 95 whenthe leaf area indexis verylargeand Table 3 compares
ratesfortropicaland temperate climatesassumingaverageratesofinsolation.Note that
theefficiencyE (g, r) fora Class I crop is about twicethevalue fora Class III crop and
thattheyieldpredictedfora Class III crop is about thesame in tropicaland temperate
climatesbecause greaterinsolationin the tropicsis balanced by smallervalues of Ed
and Sr*
In this analysis,the interception factorei emergesas a major discriminant of dry
matterproductionaccounting,on the one hand, for differences of productivity under
differentconditionsofclimateand management and,on theother,fordifferencesbetween
themean and maximumratesof productionwithina particularstand.
Secondaryproduction
Analysisof crop growthin termsof energyconversioncan readilybe extendedto
includetheeconomicyieldof a crop,secondaryproductivity, and theenergyavailableas
humanfood. The fractionof usefulenergyremovedfroma crop can be writtenEu and
will assume differentvalues dependingon whetherthe materialis harvestedby man-
in whichcase EUwillbe almostthesame as thecropfactordiscussedbyHolliday(1973)-
or grazedby animals.For example,in Queen ElizabethPark,Uganda, grazingby large
herbivoresremovesabout one-thirdof thevegetationproducedannually(C. D. Pigott,
personalcommunication). Assumingthathalftheenergyof thismaterialcan be meta-
bolizedE,,-0 15. At a levelof 500 g m2 year-', theannualproductionofdrymatteris
equivalentto 0-27W m-2 or to E (g, r) = 2 x 10-4. (Referring to Table 2, thisefficiency
can be derivedbyputtingei = 0-2in thecolumnforsubsistenceagriculture.) The power
availableto herbivores 0 15 x 0-27W m-2 and fortheMweyaPeninsulawith
is therefore
an area of 5 km-2 thepoweris 200 kW. Accordingto Eltringham (1973) thetotalmeta-
bolismof all the herbivoresin thisarea is 252 x 106 cal h-' whichis equivalentto 290
kW in good agreementwiththepowerequivalentof consumeddrymatter.
As the metabolismof large animalsis nearlyproportionalto theirsurfacearea, it is
possibleto calculatethetotalskinsurfacethata givenweightof vegetationcan support.
The rateof fastingmetabolismis about 40-50 W m-2 and allowingfortheextraenergy
used in movementand in digestion,a roundfigureof 60 W m-2 will be adopted here.
Dividingtheenergyavailable fromdry matterin W m-2 by 60 W m-2 givesa skin
area index,thearea of herbivoresupportedby unitarea of ground.At Queen Elizabeth
Park wheretheenergysupplyis 4 x 10-2 W m-2, the skinarea indexis 4 x 10-2/60or
7 x 10-4, i.e. 1500m2 of groundarea are neededto produceenoughenergyfor1 m2 of
animal.The same ratiocan be derivedby dividing1360W m-2 by 60 to geta skinarea
indexcorresponding to thesolarconstant;thisis 22-7.Multiplying 22-7bytheefficiency
E,E (g, r) = x the
3 10-5 gives figure of 7 x to
10-4 appropriate Queen ElizabethPark.
To estimatetheenergyavailablefroma crop as humanfood,it maybe assumedthat
the economicyieldis a fixedfractionof totaldrymatterproductionand thata further
fractionEf is retainedafterlosses in harvesting, storageand preparationforthe table.
G
764 Productivity
AssumingEU = 0 5 and ES = 0 5, the efficiency withwhichfood energyis producedin
subsistenceagricultureis 0-25x 0 5 x 10-4 or 1P3x 10-5 usingthefactorsin Table 2.
On a basis of 60 W m-2, a man witha skinarea of 1P7m2willmetabolizeenergyat a
mean rate of 100 W (2100 cal day-1) consistentwithfiguresof energyintakein the
tropics(but 50% smallerthan the average intakein developedcountries).The solar
constantof 1360 W m-2 is equivalentto the energyproduced by 13-6men m-2 or
13 600 menha-1. In regionssupportedbysubsistence wherea relatively
agriculture small
fractionoffood energyis derivedfrommeatand animalproducts,thenumberof people
per unitarea of cultivatedland can be predictedby multiplying thepersonalequivalent
ofthesolarconstantbytheefficiency ES s,4E (g, r). For example,givena levelofsubsistence
agriculturewithEs = 0 05 theequivalentpopulationis 1P36x 105x 1P3x 10- 5 or 1P8men
ha-1, and sincehalfa hectareis about thelimitthatone man can farmwithtraditional
tools (Boshoff1973),subsistencefarmingat thislevelmustbe close to thelowerlimitat
whichcommunities can survive.Generalizing,Fig. 15 showsthe predictedrelationship
4 I005 0; 10 TropicalS. America
S.E. Asia
-r / -Middle America
3_ - Middle S. Asia
3
W. Africa
N. Africa
S. Europe
S 2 - E. Africa
W / - S.W Asia
o I Middle Africa
5 10
E (Or) X104
FIG. 15. Estimateof populationdensitiesin relationto interception

efficiency
8g and
ofcropproduction
efficiency e (g, r).
betweenpopulationper unitarea of cultivatedland,thecombinedefficiency E (g, r) and

theinterception efficiency
ei using theparameters in Table 2 with8U8f= 0-25.The same
figureshowstheactualpopulationdensitiesin underdeveloped countriesabstractedfrom
UnitedNationsstatistics by Revelle(1966). Thesedensitiesare consistentwitha rangeof
values of E (g, r) from3 to 12 x 10-4 or of ei from0-03to 0412.
CONCLUSIONS
(1) The efficiencywithwhichplantsstoresolarenergycan be expressedas theproduct
of seven factorswhichdescribethe dependenceof drymatterproductionon latitude
and season (Eg), on cloudinessand the aerosol contentof the atmosphere(Sa), on the
spectralcompositionofradiation(Es) and thequantumneedofthephotochemical process
(sq), on leaf area index and leaf arrangement (8k), on the of
concentration CO2 in the
canopyand thediffusion resistanceofindividualleaves(Ed),and on thefractionof assimi-
latesused in respiration(er). The analysiscan be extended to estimateanimaland human
populationsfromtherelationbetweenprimaryproductionand ratesof metabolism.
(2) Estimatesof drymatterproductionfora rangeof agricultural crops growingbe-
J. L. MONTEITH 765
tween140 S and 430 N agreewellwithmeasurements duringthevegetative stageofgrowth
but overestimate productionduringreproductive growth,eitherbecause photosynthesis
ratesare overestimated or respirationratesare underestimated or both. The agreement
for vegetativegrowthis not necessarilypeculiar to one particularmodel of canopy
photosynthesis but is probablycommonto severalmodels.
(3) The main limitationsof the modelscurrently used to predictcrop growthfrom
photosynthesisratesare: (i) ignoranceabout whetherthedecreaseof Ed as a leafages is a
resultmainlyof an increasein theresistanceto CO2 diffusion withinmesophylltissueor
of a decreasein the concentration of carboxylating enzymes;(ii) ignoranceabout the
wayin whichratesof respiration are relatedto thecontemporary rateof photosynthesis
and to theaccumulateddryweightof thestand; and (iii) ignoranceabout waysin which
the partitioning of assimilatesmay be determinedby an interactionof environmental
factorsand endogenouscontrols.The needto use measurements insteadofpredictionsof
leaf area index is a weaknessof most models. The ELCROS model developedby de
Wit & Brouwer(1970) includesan estimateof the rate at whichleaves expand after
germination but muchmoreresearchis neededbothin thefieldand in thegrowthroom
to determine howleafexpansionratesdependon environmental factorssuchas tempera-
tureand theavailabilityof waterand nutrients in the root zone.
ACKNOWLEDGMENTS
I am verygratefulto mycolleaguesMr K. Gregsonand Dr M. H. Unsworthwho wrote
computerprogramsto calculate photosynthesis rates and insolationrespectively;to
Dr J. Kowal, Instituteof AgriculturalResearch, Samaru and Dr F. J. Wangati,
E.A.A.F.R.O., Nairobi,forrecordsof solar radiation;and to Dr J. Elston of Reading
Universityformanystimulating discussionson the subjectof thispaper.
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GPP in Tropical Ecosystems (Monteith, 1972)

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GPP in Tropical Ecosystems (Monteith, 1972)

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Solar Radiation and Productivity in Tropical Ecosystems

In thermodynamic terms,ecosystemsare machinessuppliedwithenergyfroman ex-

cloud or of particulatesand to expressthisfluxas a fractionof theextraterrestrial

"_, > > Aerosol

Chlorophyllabsorbsverystrongly in theblue and redregionsof thespectrumso that

Sq = 10 x (3-6x l0- 19) = 0-215.

FIG. 6. Carbondioxideexchangeof isolatedleafas function EA represents

Two limitshave now been establishedfor rates of photosynthesis by singleleaves:

_S[Ao{(l-s)a+b/I} '+A1{(l-s)a+b/'I}-'] dt (4)

In thelimitwhenthe leaf area indexis verylarge so thatthe canopyintercepts all the

Whentheradiation i) b/Iis muchlargerthan(1 -s)a andit

Assuminga Q10 of 2 and dividingthroughby P therelationbecomes

Br= 1- {00625+O000375 WIP}2T/1,

TESTS OF THE ANALYSIS

I00 200 300 100 200

100 200 100 1OO

GENERALIZATION AND EXTENSIONS

Table 3. Maximumratesof drymatterproduction

Differences of latitudeand cloudinessare takenintoaccountby Eg and sa respectively.

4 I005 0; 10 TropicalS. America

FIG. 15. Estimateof populationdensitiesin relationto interception

betweenpopulationper unitarea of cultivatedland,thecombinedefficiency E (g, r) and

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