Royal Swedish Academy of Sciences

Human Appropriation of Net Primary Production as an Environmental Indicator: Implications

for Sustainable Development
Author(s): Helmut Haberl
Source: Ambio, Vol. 26, No. 3 (May, 1997), pp. 143-146
Published by: Springer on behalf of Royal Swedish Academy of Sciences
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 Report HelmutHaberl
Human Appropriation of Net Primary
Production as An Environmental Indicator:
Implications for Sustainable Development
The human appropriation of net primary production (NPP)
significantly alters the energy flow of ecosystems. The
NPP-appropriation, defined as the difference between the
NPP of the hypothetical undisturbed vegetation and the
amount of biomass currently available in ecological cycles,
is investigated for the 99 political districts of Austria (1990).
Calculations are based on data for land-use, forestry, yield,
and climate. Total aboveground NPP of the actual vegeta-
tion was found to be 7% less than that of the potential
naturalvegetation. Additionally,34% of potential production
is harvested, resulting in a total reduction of ecologically
available aboveground NPP of 41 %. Since this could have
significant ecological effects, e.g. on biodiversity, it is of
potential interest for strategies of sustainable development,
indicators for stresses on the environment, and the
environmental effects of increased utilization of biomass.
INTRODUCTION
The productionof biomass by green plants is the main energetic
basis of life on earth and provides the primaryinput for most
food chainsof all types (herbivory,carnivory,detritivory).While
hunters and gatherersdwelled upon the products of photosyn-
thesis much like any other kind of animal species, thus, reach-
ing only very small densities, the culturalevolution of human-
ity has seen a tremendousintensificationof biomassuse (1). This
could only be achieved by a transformationof naturalecosys-
tems into managed ones with an increasing numberof ecosys-
tem variables being controlled. While agricultureallowed the
harvest of more biomass per unit area, it did not generally in-
crease the average productivityin terms of total energy or car-
bon fixation. Additionally,ever-increasingareasare used for the
constructionof buildings,roads, etc. and thus theirprimarypro-
ductivity is reduced.
While ecologists were concerned about the possible exhaus-
tion of the world's biomass resources alreadyvery early (2), it
was the seminalpaperof Vitousek et al. (3) thatopened the door
for a broaderdiscussion of the problem. In the meantime, it is
estimatedthat the human appropriationof the productsof pho-
tosynthesisamountsto between 25 and 39% of the global terres-
trial net primaryproduction(NPP) (3, 4). This result inspired
Meadows et al. to worryaboutthe possible consequencesof the
humanworld biomass use, which can be expected as a result of
the projected doubling of the world population and economy
within the next 20 or 30 years (5) and was broadly noticed in
the discussion on sustainabledevelopment (6, 7). This discus-
sion revealed that NPP is an importantlimiting resourcefor the
future development of humanityand currentlevels of NPP ap-
propriationalreadyappearto be considerable(3). Moreover,cur-
rent strategies of sustainabledevelopment in the energy sector
partly rely on the substitutionof fossil fuels by biomass in or-
der to reduce CO2emissions-a strategythat would furtherin-
crease NPP-appropriation.
Ambio Vol. 26 No. 3, May 1997 ? Royal Swedish Academy of Sciences 1997
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This article relies on a spatiallyhighly resolved study on the
societal NPP appropriationin Austria. Austria is an industrial-
ized countrywith mediumpopulationdensity (area83 000 km2,
population7.8 million). Forestscover about45% of its area.This
is a ratherhigh percentagefor CentralEuropeanstandardsdue
to the mountainouslandscape.
MATERIALSAND METHODS
In this article,NPP appropriationis defined as the differencebe-
tween the NPP of the potentialnaturalvegetation(NPPO,i.e. the
vegetationthatwould prevailif humaninterferencewere absent)
and the amountof biomass currentlyavailablein ecological cy-
cles (NPPt). Two processes contribute to NPP appropriation
(NPPa):i) changes in the averageproductivity(NPP per unit area
and year) of ecosystems, e.g. the construction of a road in a
forestedecosystem; and ii) harvest.If the NPP of the actualveg-
etation is denoted as NPPactand harvestas NPPh,total NPP ap-
propriationcan be calculatedwith the formula:
NPPa= NPPo- NPPtwith NPPt= NPPaCt
- NPPh
All calculationswere performedon the spatial level of com-
munities(Austriaconsists of approximately2350 communities).
Since therearefew reliabledataon the subterranean primarypro-
ductionof forest ecosystems (it was significantlyunderestimated
in previousresearch(8-10)), this articlefocuses on aboveground
net primary production (ANPP) which can be assessed with
greateraccuracy.(see Haberl(11) for an in-depthdescriptionof
all appliedmethods).
The ANPP of the potentialnaturalvegetation in Austriawas
estimated with two independentmethods: i) The average pro-
ductivity of different types of naturalvegetation was assessed
on the basis of the literatureavailable (11), applyingregression
analyses on the relationbetween mean annualtemperature,pre-
cipitation and net productivityin forest ecosystems using data
compiled by Cannel (12) and the climate data of Walter and
Lieth (13); ii) The so-calledMiamimodell of Lieth (14) was used
with a correctionfor Lieth's assumptionon subterraneanNPP.
Data on mean annual precipitationand long-term temperature
averages were obtainedfrom the Austrianassociation of mete-
orology (15) and modified to reflect elevation. The areaof each
communitywas distributedover six elevation classes (less than
600 m, 600-1300 m, 1300-1700 m, 1700-2200 m, 2200-2600
m, more than2600 m) assessed by a geographicinformationsys-
tem (Loibl, pers. comm.) which served to determinethe poten-
tial natural vegetation. The data used for the calculation of
ANPPo and ANPPaCt in naturalecosystems (and actual forests,
as describedbelow) are given in Table 1.
The productivityof the actual vegetation was calculated by
using land-use data as assessed by the AustrianCentralStatisti-
cal Office (16). The productivityof agriculturalareas(crops and
meadows) was estimated by using harvest factors of the form
NPP = H x F, where H is the commercialharvest and F an ap-
propriatefactor for total or abovegroundproductivity.Harvest
143
factors were taken from the literature(14, ...

.......
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.. .... "Ill- ..........

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I'll,
................ ... ...... . .........
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17-20). The productivityof grazingland was . .......... . . . ...... ..

........ ...... ...... . ........ ..

......

..........
. . ................
..........
.. ...................
.....

'NM
calculated from average productivity esti- Mi
ttt
. . ...... .............
. . .... ... . ... ... ... ... . . . . . .

mates, dependingon elevation, based on the ........... ...

.... .....
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.......... iW
... nn

EJ J g?n!F
......................... ... !MTi

literature. The productivity of forests was ... . . ....... ....

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HHE:
...........
.............

tr
...........

estimatedby two independentmethods,first ..................

..............
.
. .......
..... ..... .

by harvest factors from the literature(11), .. ..........

. ........

using the Austrianforest inventory(21), and

........... .
... ....... .. .......... ............ . ........................ ....

....... ...... . .....

:.::.nf,l-j .....

second by assuming the average productiv- ......

.. ...... . ...........
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E . .... .
4,V
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ity by forest type, modified by elevation ..... ..... ......

........ ....... ......
... ... ... s-ss
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.. ....... .
ss;-N ..........
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. . .......... ......I", ::: -s, s::::::::: .::. fg
NM ........... .......... . . .......
....... ......... ........................ .. ..................................................... ............. ............

class (Table 1). .. .... . .. ... ... .... ... .. . . . ..... . ...... .......... . N. . :":.. . - .1-11.1- ... . . ..... ............. ....

Harvestwas calculatedfrom Austrianag-

riculturaland forestry statistics (22, 23). Subterraneanparts of Together with a straightforwardestimation of harvest (512
crops like potatoes (about 1% of total harvest) were counted as PJ yri- or 27.6 mill. t yr-' DM) ANPP appropriation(ANPPa)
"above-ground". Agriculturalbiomasswas convertedto dry mass can be assessed to amountto 617 PJ yrf- or 41.1% of potential
and calorific value using standardtables on nutritive value of abovegroundproductionin Austria(Fig. 1).
the materialsunderconsideration(24, 25), wood was treatedin My tentative calculations on total NPP and its appropriation
the same way on the basis of tables on species-specific dry-mat- show that the difference between NPPo and NPP,actshould be
ter content and calorific value. much higher than that for aboveground NPP, because the
belowgroundproductivityof forests is considerablyhigher than
that of annualcrops; even in cases where abovegroundproduc-
RESULTS tivity is similaror crops are more productive.Total NPP appro-
Both calculationmethods for the ANPPOof the hypotheticalun- priation,however, is smaller,since mainly abovegroundbiomass
disturbedvegetation of Austria led to almost the same result. is harvested.As these results are ratheruncertain,they will not
While Lieth's Miami model predicts the ANPP to be 1445 be discussed here in detail (11).
PJ yr-1(74 mill. t yr'- dry matter,DM), the assumptionof aver- As Figure 2 shows, there is more NPP appropriationin dis-
age productivitiesdependent on elevation gave an estimate of tricts with high ANPPOthan in districtswith low ANPPO.Obvi-
1501 PJ yr71(77.6 mill. t yr-' DM). Since Lieth'smodel is based ously, fertileregions aremore intensivelycultivatedand a higher
on ratherold productivitydata which tended to underestimate share of their net primaryproductionis harvested. Settlements
productivity(26), the higher value is believed to be more reli- and roads are also preferably situated in low, fertile regions.
able and taken as a referencepoint. The averageproductivityof Thus, the higher the level of ANPPO,the higher the proportion
the Austrianvegetation is 0.93 kg m-2yr' or 17.9 MJ m-2 yr-'. of appropriatedANPP. Naturalfertilityexplains 50% of the vari-
Humanactivities, above all agricultureand construction,have ance of ANPPJANPPO,this relation being significant at p <
significantly lowered the productivityof the vegetation in Aus- 0.001 (chi square).
tria. ANPPaCt was estimated to be 1396 PJ yr-1which is 105 PJ
yri' (7.6%) lower than the ANPPO.In terms of dry matter,the
difference is smaller (ANPPaCt= 74.2 mill. t yr'- DM), since the DISCUSSION
calorific value of forest biomass is higher than that of most cul- The reliability of the results can be assumed to be high for
tivated herbaceousplants. About 50 PJ yf-' of this reductionis abovegroundNPP. A previousstudy (27), which used much sim-
due to construction.The difference between the two independ- pler calculationmethodsand a much narrowerdatabase, yielded
ent methods used to estimate primaryproductionof forests (el- rather similar results. The results on average productivity of
evation classes and forest inventory) was only about 2%, with ANPPOand ANPPaCt are well in line with the comparabledata
forest inventories giving a slightly higher value. The value re- in recent large-scale productivitystudies (14, 28-30). The ap-
ported above is based on elevation classes. It should be men- propriationof ANPP in Austria is higher than the estimates for
tioned that these estimates are conservative, since the statistical global NPP appropriation,rangingfrom 25 to 39% (3, 4). Nev-
data on land-use for constructionare believed to be lower than ertheless, it can be assumed that values for other highly indus-
actualvalues and many of the underlyingassumptions(11) were trialized western and CentralEuropeancountries may be even
made with great caution to avoid a possible overestimationof higher: They typically have less than the Austrian45% of for-
NPP appropriation. ests with rather low NPP appropriationper M2. Furthermore,

_ te F_ _
[PJ/yFJ rpJ~~r-1l 1.5011
.........~~~~~~..................... -- ----- ...
----_
-----------------
.
1.400 . . __j - - - i----.-- ........................
1.396 --- - - - - . ~----- -oo i;i
--_-----

ANPPA I
1.200
0NPPh 617

1.000 (4,1%) 10.

ANPPo _ ct_,
ANPP_a........ .NPP -------------------------- 4

0
400_ 88% ._

Figure 2. NPP appropriation greatly reduces the spatial differences

between the energy Input of different terrestrial ecosystem types. While
ANPPo ANPPact
ANPPOand ANPP.CJ fall within a range between 8 and 22 MJ m-2yr', the
Figure 1. Appropriation of aboveground net primary production (ANPP) amount of energy actually remaining serving as Input for the
in Austria 1990. Of the 1501 PJ yr-' which would be available in natural heterotrophic food chains Is much more evenly distributed and ranges
ecosystems If human interference were absent, only 884 PJ yr1- can from about 7 to 15 MJ m 2 yr-'. ANPPOand ANPPC,are the aboveground
actually serve as energy Input of all heterotrophic food chains. NPP of the potential vegetation and the actual vegetation, respectively.

144 i Royal Swedish Academy of Sciences 1997 Ambio Vol. 26 No. 3, May 1997

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12.6%of the Austrianterritoryis above 1800 m elevation,where
no NPP appropriationwas assumed to occur. In other countries
agriculturalareas,roadsand buildingsare likely to cover a much
higherpercentageof the surface.
These results raise the question of which effects humanNPP
appropriationmay have on naturalecosystems. Obviously,it sig-
nificantly alters the energy flow of naturalecosystems and thus
may be seen as an indicatorfor the intensityof humaninterven-
tions into natural ecosystem processes (27). But what do we
know aboutits likely effects on the structureand functioningof
ecosystems?
If we follow the argumentsof Hutchinsonin his famous pa-
per "Homageto SantaRosalia, or why are there so many kinds
of animals?"(31), we may suspect that a reduction of energy
flow is likely to cause a reductionof the length of food chains.
His argumentwas that since less than 10%of the energy avail-
able at the level n of a food chain can be gatheredat the level
n + 1, food chains cannot be very long.
Although it appearsto be clear that the length of food chains
is ultimately constrainedby the amount of energy available, it
may well be that in many cases other factors-body size, stabi-
lity of food chains, etc.-may be more important.Until now,
empirical studies failed to produce unequivocal results on this
matter.While Briand and Cohen found no correlationbetween
energyflow and food chain lengthin an analysisof 38 food webs
(32), Yodzis could show that ectothermfood chains are signifi-
cantly longerthanendothermfood chains (33). Since ectotherms
convertfood to secondaryproductionmuch more efficientlythan
endothermsthis result supportsthe energy theory of food chain
length regulation. In addition, only recently an experimental
study has shown the importanceof energy availability on food
chain length (34). In model calculations, Oksanen showed that
the amountof energy available per unit area greatly influences
food chain structure,as far as vertebratesare concerned: Ac-
cording to the results of his models, big grazers will dominate
ecosystems with an average aboveground productivity below
approximately 12 MJ m-2 yr-', while complex food chains in
which the herbivoresareregulatedby carnivores,andhave much
lower populationdensity, prevail in richerhabitats(35).
If it is true that a reductionof energy flow reduces the length
of food chains, then a second assertionof Hutchinson(31) may
also prove correct, namely that the amount of energy available
exerts an importantinfluenceon species diversity.In the last two
decades this idea experienceda renaissanceas the so-called spe-
cies-energy theory of biodiversity (4, 36-38). In short, the spe-
cies-energy theorypredictsthatthe numberof species which can
inhabit a certain environmentincreases with the amount of en-
ergy available;conversely, the numberof species will decrease,
if energy flow is reduced (Fig. 3). The rationale behind this
- - - - - - - -
-
N N1 -_
number
Of
species N2 - - -
E2 El
E energy flow (e.g. NPP)
Figure3. Species-energy curves demonstratethe relationbetween
energy flow (E)and species richness (N).Ifenergy flow is reduced,
species-energy theory predictsa reductionof species richness.
Ambio Vol. 26 No. 3, May 1997 ? Royal Swedish Academy of Sciences 1997
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theory is that in habitatswith abundantresources rivaling spe-
cies will be able to specialize with respectto more gradientsand
thus can avoid extinction due to Gauses principle of competi-
tion exclusion (36). While in poor habitats there are few,
generalistic species, in resource-richhabitats many specialists
prevail. One reason for this are the "costs of commonness",i.e.
negative effects of high population densities, e.g. parasitism,
pests, specialized predators(38).
The species-energytheory has been shown to be an extension
of species-areatheory (37), which relies on the theoryof island-
biogeographyby MacArthurand Wilson (39). The core of this
theoryis thatthe numberof species on an island is a steady state
between immigration(and speciation) and extinction, and the
bigger the island, the greatera numberof species it is able to
support.Species-energytheory claims that this assertioncan be
explained by the fact that ceteris paribus bigger islands provide
more energy, and predicts that among islands of the same size
more productive ones will supporta higher numberof species
(4). The species-energy theory is not only able to explain the
gradientof species diversity from the poles to the equator,but
has also been empiricallytested and verified (38, 40-42).
Even if the species-energy theory may be an innovative ap-
proachin biodiversityresearch,it is currentlynot generally ac-
cepted. For example, it cannot explain the "paradoxof enrich-
ment"(43); i.e. the observationthatnutrient-rich(and thus more
productive)habitatsmay have lower species diversity than less
fertilized ones, a phenomenon Tilman has explained with a
microeconomicmodel (44). Since such a model fails to explain
the big biogeographicalbiodiversity gradientfrom the poles to
the equator,however, it cannot claim to be the uniquetheoryof
biodiversity. In general, it is likely that the explanation of
biodiversitypatternsrequiresmore thanone scientific approach.
As far as the Austriandata are concerned,my results are con-
sistent with the species-energy theory. If the propertiesof the
species-energycurve of Wright(4) are assumed, species-energy
theory predicts that in Austria between 5 and 13% of the spe-
cies should have gone extinct up to now. Actual surveys show
that 8% of the bird species, 7-14% of the reptiles (but no am-
phibians)have gone extinct in Austria(45, 46). However, since
this may just be coincidental,it is not a very strongargumentin
favor of the species-energytheory. But it does ascertainthatthe
theory does not contradictthe data. Furtherresearch,based on
the datanow available,which have a high spatialresolution,will
be directed towards an attemptto explain biodiversitypatterns
with variationsof availableenergy (ANPP,).
CONCLUSIONSREGARDINGSUSTAINABLE
DEVELOPMENT
Even if the effect of NPP appropriationon biodiversity is yet
unproven,it is obvious that the currentlevel of NPP appropria-
tion constitutesa significantinterventioninto the naturalenergy
flow of ecosystems. The potential effects this interferencemay
have, and which are currentlynot well understood,demandthat
NPP appropriationshould be regardedas an importantindica-
tor for pressureson the environment.Since the NPP of a natu-
ral ecosystem appearsto be an insurmountablelimit (globally
as well as at the local level), this indicatorshould be considered
as a core parameterfor sustainabledevelopment(47).
Many strategiesfor sustainabledevelopmentin the energysec-
tor seek to promotethe substitutionof fossil fuels with biomass
(48). Most of these strategies, however, imply an increase of
biomass harvest and thus are likely to contributeto an increase
of NPP appropriation,e.g. an increase of firewood combustion
for heating purposes,and thus could threatenbiodiversity.As a
consequence, targetconflicts may exist between C02-reduction
and the conservationof biodiversity, which are both important
aspects of sustainabledevelopment.Moreover,the calculations
145
presented above show that the idea of a simple substitutionof
fossil fuels with biomass would not work, at least in Austria,
simply because the total energy input of 1680 PJ yr-f (including
biomass for nutrition) of Austria already is higher than the
ANPPact(1396 PJ yr-F)(49).
Which strategiesmay be envisagedto avoid this potentialcon-
flict? One possible part of a solution is the cascade use of
biomass. This means that waste biomass should be used for en-
ergy generationinsteadof harvestingmore biomass. This would
permit an increase of biomass use for energy generationwith-
out augmentingNPP appropriation.One example is the produc-
tion of biogas from wet biomass waste, e.g. animal manure.
Potentials for the generation of additional energy from used
biomass should be systematicallyinvestigated.
Anotherstrategy,which has the advantageof alleviatingother
ecological problemsas well, is based on the observationthatthe
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146 ? Royal Swedish Academy of Sciences 1997
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50. For help, advice, anddiscussionsI am indebtedto W. Bittermann,M. Fischer-Kowalski,
B. Hammer,W. Huttler,W. Loibl, H. Payer, H. Schandl, V. Winiwarter,H. Zangerl-
Weisz, and two anonymousreferees.
51. First submitted28 November 1995. Accepted for publication after revision 19 June
1996.
Helmut Haberl,PhD, is currentlywith the Austrian Institute
of Applied Ecological Research and at the Interdisciplinary
Institute of Research and Continuing Education (IFF),
Departmentof Social Ecology. He works on energy and the
environment, environmental indicators, societal metabolism
and the colonization of nature, and sustainable
development. His address: IFF,Social Ecology,
P.O. Box 232, Seidengasse 13, A-1070 Vienna, Austria,
e-mail: helmut.haberl@univie.ac.at
Ambio Vol. 26 No. 3, May 1997