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l
Human Alrerarion of rhe
Clobal Nirrogen Cycle:
Causes and Consequences
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SbMMAkY
Human acriviries are grearly increasing rhe amounr of nirrogen cycling berween rhe living world and rhe soil, warer, and
armosphere. |n facr, humans have already doubled rhe rare of nirrogen enrering rhe landbased nirrogen cycle, and rhar
rare is conrinuing ro climb. This humandriven global change is having serious impacrs on ecosysrems around rhe world
because nirrogen is essenrial ro living organisms and irs availabiliry plays a crucial role in rhe organizarion and funcrioning
of rhe world`s ecosysrems. |n many ecosysrems on land and sea, rhe supply of nirrogen is a key facror conrrolling rhe
narure and diversiry of planr life, rhe popularion dynamics of borh grazing animals and rheir predarors, and viral ecologi
cal processes such as planr producriviry and rhe cycling of carbon and soil minerals. This is rrue nor only in wild or
unmanaged sysrems bur in mosr croplands and foresrry planrarions as well. Excessive nirrogen addirions can pollure
ecosysrems and alrer borh rheir ecological funcrioning and rhe living communiries rhey supporr.
Mosr of rhe human acriviries responsible for rhe increase in global nirrogen are local in scale, from rhe producrion and use
of nirrogen ferrilizers ro rhe burning of fossil fuels in auromobiles, power generarion planrs, and indusrries. However,
human acriviries have nor only increased rhe supply bur enhanced rhe global movemenr of various forms of nirrogen
rhrough air and warer. because of rhis increased mobiliry, excess nirrogen from human acriviries has serious and long
rerm environmenral consequences for large regions of rhe Earrh.
The impacrs of human dominarion of rhe nirrogen cycle rhar we have idenrified wirh cerrainry include:
|ncreased global concenrrarions of nirrous oxide (N
J
Oj, a porenr greenhouse gas, in rhe armosphere as
well as increased regional concenrrarions of orher oxides of nirrogen (including nirric oxide, NOj rhar
drive rhe formarion of phorochemical smog:
Losses of soil nurrienrs such as calcium and porassium rhar are essenrial for longrerm soil ferriliry:
Subsranrial acidificarion of soils and of rhe warers of srreams and lakes in several regions:
Crearly increased rransporr of nirrogen by rivers inro esruaries and coasral warers where ir is a major
polluranr.
ve are also confidenr rhar human alrerarions of rhe nirrogen cycle have:
Accelerared losses of biological diversiry, especially among planrs adapred ro lownirrogen soils, and
subsequenrly, rhe animals and microbes rhar depend on rhese planrs:
Caused changes in rhe planr and animal life and ecological processes of esruarine and nearshore
ecosysrems, and conrribured ro longrerm declines in coasral marine fisheries.
Narional and inrernarional policies should arrempr ro reduce rhese impacrs rhrough rhe developmenr and widespread
disseminarion of more efficienr fossil fuel combusrion rechnologies and farm managemenr pracrices rhar reduce rhe
burgeoning demand for and release of nirrogenous ferrilizers.
Human A|terat|on ot the
6|oha| N|trogen Cyc|e:
Causes anJ Consequences
Issues |n |co|ogy Numher 1 Spr|ng 1997
J
INTkODbCTION
This reporr presenrs an overview of rhe currenr
scienrific undersranding of humandriven changes ro rhe
global nirrogen cycle and rheir consequences. |r also
addresses policy and managemenr oprions rhar could help
moderare rhese changes in rhe nirrogen cycle and rheir
impacrs.
TH| NITkO6|N CYCL|
Nirrogen is an essenrial componenr of proreins,
generic marerial, chlorophyll, and orher key organic mol
ecules. All organisms require nirrogen in order ro live. |r
ranks fourrh behind oxygen, carbon, and hydrogen as
rhe mosr common chemical elemenr in living rissues. Unril
human acriviries began ro alrer rhe narural cycle (|igure
1j, however, nirrogen was only scanrily available ro much
of rhe biological world. As a resulr, nirrogen served as
1
Human A|terat|on ot the 6|oha| N|trogen Cyc|e:
Causes anJ Consequences
by
Perer M. virousek, Chair, 1ohn Aber, Roberr v. Howarrh,
Cene E. Likens, Pamela A. Marson, David v. Schindler,
villiam H. Schlesinger, and C. David Tilman
one of rhe major limiring facrors rhar conrrolled rhe dy
namics, biodiversiry, and funcrioning of many ecosysrems.
The Earrh`s armosphere is S percenr nirrogen
gas, bur mosr planrs and animals cannor use nirrogen
gas direcrly from rhe air as rhey do carbon dioxide and
oxygen. |nsread, planrs and all organisms from rhe
grazing animals ro rhe predarors ro rhe decomposers rhar
ulrimarely secure rheir nourishmenr from rhe organic
marerials synrhesized by planrs musr wair for nirro
gen ro be fixed," rhar is, pulled from rhe air and bonded
ro hydrogen or oxygen ro form inorganic compounds,
mainly ammonium (NH
4
j and nirrare (NO
J
j, rhar rhey
can use.
The amounr of gaseous nirrogen being fixed ar
any given rime by narural processes represenrs only a
small addirion ro rhe pool of previously fixed nirrogen
rhar cycles among rhe living and nonliving componenrs
of rhe Earrh`s ecosysrems. Mosr of rhar nirrogen, roo,
is unavailable, locked up in soil organic marrer par
||gure 1Simplified diagram of rhe nirrogen cycle. Adapred from Eac|rcameata| Sc|eace, Third Edirion by 1onarhon
Turk and Amos Turk, &19S4 by Saunders College Publishing, reproduced by permission of rhe publisher.
J
Issues |n |co|ogy Numher 1 Spr|ng 1997
a million merric rons of nirrogen. vorldwide, lighrning,
for insrance, fixes less rhan 10 Tg of nirrogen per year
maybe even less rhan 5 Tg. Microbes are rhe major
narural suppliers of new biologically available nirrogen.
before rhe widespread planring of legume crops, rerres
rrial organisms probably fixed berween 90 and 140 Tg
of nirrogen per year. A reasonable upper bound for rhe
rare of narural nirrogen fixarion on land is rhus abour
140 Tg of N per year.
HbMAN-DkIv|N NITkO6|N |IXATION
During rhe pasr cenrury, human acriviries clearly
have accelerared rhe rare of nirrogen fixarion on land,
effecrively doubling rhe annual rransfer of nirrogen from
rhe vasr bur unavailable armospheric pool ro rhe biologi
cally available forms. The major sources of rhis enhanced
supply include indusrrial processes rhar produce nirro
gen ferrilizers, rhe combusrion of fossil fuels, and rhe
culrivarion of soybeans, peas, and orher crops rhar hosr
symbioric nirrogenfixing bacreria. |urrhermore, human
acriviry is also speeding up rhe release of nirrogen from
longrerm srorage in soils and organic marrer.
rially rorred planr and animal remains rhar musr be
decomposed by soil microbes. These microbes release
nirrogen as ammonium or nirrare, allowing ir ro be re
cycled rhrough rhe food web. The rwo major narural
sources of new nirrogen enrering rhis cycle are nirrogen
fixing organisms and lighrning.
Nirrogenfixing organisms include a relarively
small number of algae and bacreria. Many of rhem live
free in rhe soil, bur rhe mosr imporranr ones are bacreria
rhar form close symbioric relarionships wirh higher planrs.
Symbioric nirrogenfixing bacreria such as rhe Rhizobia,
for insrance, live and work in nodules on rhe roors of
peas, beans, alfalfa and orher legumes. These bacreria
manufacrure an enzyme rhar enables rhem ro converr
gaseous nirrogen direcrly inro planrusable forms.
Lighrning may also indirecrly rransform armo
spheric nirrogen inro nirrares, which rain onro soil.
Ouanrifying rhe rare of narural nirrogen fixarion
prior ro human alrerarions of rhe cycle is difficulr bur
necessary for evaluaring rhe impacrs of humandriven
changes ro rhe global cycling of nirrogen. The srandard
unir of measuremenr for analyzing rhe global nirrogen
cycle is rhe reragram (abbreviared Tgj, which is equal ro
4
||gure 2Nirrogen is rhe major facror limiring many rerresrrial ecosysrems, including mosr of rhose in rhe
remperare zone, such as rhis oak savannah. The number and idenriries of rhe planr and animal species rhar
live in such rerresrrial ecosysrems, and rhe funcrioning of rhe ecosysrem, depends on rhe rare of nirrogen
supply ro rhe ecosysrem.
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Issues |n |co|ogy Numher 1 Spr|ng 1997
5
N|trcqca Fcrt|||zcr
|ndusrrial fixarion of nirrogen for use as ferrilizer
currenrly rorals approximarely S0 Tg per year and repre
senrs by far rhe largesr human conrriburion of new nirro
gen ro rhe global cycle (|igure Jj. Thar figure does nor
include manures and orher organic nirrogen ferrilizers,
which represenr a rransfer of alreadyfixed nirrogen from
one place ro anorher rarher rhan new fixarion.
The process of manufacruring ferrilizer by indus
rrial nirrogen fixarion was firsr developed in Cermany
during vorld var |, and ferrilizer producrion has grown
exponenrially since rhe 1940s. |n recenr years, rhe in
creasing pace of producrion and use has been rruly phe
nomenal. The amounr of indusrrially fixed nirrogen ap
plied ro crops during rhe decade from 19S0 ro 1990
more rhan equaled all indusrrial ferrilizer applied previ
ously in human hisrory.
Unril rhe lare 190s, mosr indusrrially produced
ferrilizer was applied in developed counrries. Use in rhese
regions has now srabilized while ferrilizer applicarions in
developing counrries have risen dramarically. The mo
menrum of human popularion growrh and increasing ur
banizarion ensures rhar indusrrial ferrilizer producrion will
conrinue ar high and likely acceleraring rares for decades
in order ro meer rhe escalaring demand for food.
N|trcqcaF|x|aq Crcps
Nearly one rhird of rhe Earrh`s land surface is
devored ro agriculrural and pasroral uses, and humans
have replaced large areas of diverse narural vegerarion
wirh monoculrures of soybeans, peas, alfalfa, and orher
leguminous crops and forages. because rhese planrs sup
porr symbioric nirrogenfixers, rhey derive much of rheir
nirrogen direcrly from rhe armosphere and grearly in
crease rhe rare of nirrogen fixarion previously occurring
on rhose lands. Subsranrial levels of nirrogen fixarion
also occur during culrivarion of some nonlegumes, nora
bly rice. All of rhis represenrs new, humangenerared
srocks of biologically available nirrogen. The quanriry of
nirrogen fixed by crops is more difficulr ro analyze rhan
indusrrial nirrogen producrion. Esrimares range from JJ
ro 5J Tg per year. As an average, 40 Tg will be used
here.
Fcss|| Fac| 8ara|aq
The burning of fossil fuels such as coal and oil
releases previously fixed nirrogen from longrerm sror
age in geological formarions back ro rhe armosphere in
rhe form of nirrogenbased rrace gases such as nirric
oxide. Highremperarure combusrion also fixes a small
amounr of armospheric nirrogen direcrly. Alrogerher, rhe
operarions of auromobiles, facrories, power planrs, and
orher combusrion processes emir more rhan J0 Tg per
year of fixed nirrogen ro rhe armosphere. All of ir is
rreared here as newly fixed nirrogen because ir has been
locked up for millions of years and would remain locked
up indefinirely if nor released by human acrion.
Mcb|||zat|ca c[ Stcrcd N|trcqca
besides enhancing fixarion and releasing nirro
gen from geological reservoirs, human acriviries also lib
erare nirrogen from longrerm biological srorage pools
such as soil organic marrer and rree rrunks, conrriburing
furrher ro rhe proliferarion of biologically available nirro
gen. These acriviries include rhe burning of foresrs, wood
fuels, and grasslands, which emirs more rhan 40 Tg per
||gure 3The pace of many hu
mancaused global changes has
increased srarkly in modern his
rory, bur none so rapidly as in
dusrrial producrion of nirrogen
ferrilizer, which has grown ex
ponenrially since rhe 1940s.
The charr shows rhe year which
changes in human popularion
growrh, carbon dioxide release,
deforesrarion, and ferrilizer pro
ducrion had reached J5%,
50%, and 5% of rhe exrenr
seen in rhe lare 19S0s. Revised
from Kares er al. (1990j.
25%
50%
75%
1700 1800 1900 1975
Deforestation
CO release
2
Human population
Industrial N fertilizer
Sources of Human-Caused Alteration to the Global Nitrogen Cycle
Issues |n |co|ogy Numher 1 Spr|ng 1997
year of nirrogen: rhe draining of werlands, which sers
rhe srage for oxidarion of soil organic marrer rhar could
mobilize 10 Tg per year or more of nirrogen: and land
clearing for crops, which could mobilize J0 Tg per year
from soils.
There are subsranrial scienrific uncerrainries abour
borh rhe quanriry and rhe fare of nirrogen mobilized by
such acriviries. Taken rogerher, however, rhey could con
rribure significanrly ro changes in rhe global nirrogen
cycle.
Hamaa \crsas Natara| N|trcqca F|xat|ca
Overall, ferrilizer producrion, legume crops, and
fossil fuel burning deposir approximarely 140 Tg of new
nirrogen inro landbased ecosysrems each year, a figure
rhar equals rhe upper esrimares for nirrogen fixed naru
rally by organisms in rhese ecosysrems. Orher human
acriviries liberare and make available half again rhar much
nirrogen. |rom rhis evidence, ir is fair ro conclude rhar
human acriviries have ar leasr doubled rhe rransfer of
nirrogen from rhe armosphere inro rhe landbased bio
logical nirrogen cycle.
This exrra nirrogen is spread unevenly across rhe
Earrh`s surface: Some areas such as norrhern Europe are
being alrered profoundly while orhers such as remore re
gions in rhe Sourhern Hemisphere receive lirrle direcr in
pur of humangenerared nirrogen. Yer no region remains
unaffecred. The increase in fixed nirrogen circularing
around rhe globe and falling ro rhe ground as wer or dry
deposirion is readily derecrable, even in cores drilled from
rhe glacial ice of Creenland.
IMPACTS ON TH| ATMOSPH|k|
One major consequence of humandriven alrer
arions in rhe nirrogen cycle has been regional and global
change in rhe chemisrry of rhe armosphere (|igure 4j
specifically, increased emissions of nirrogenbased rrace
gases such as nirrous oxide, nirric oxide, and ammonia
(NH
J
j. Alrhough such releases have received less arren
rion rhan increased emissions of carbon dioxide and vari
ous sulfur compounds, rhe rrace nirrogen gases cause
environmenral effecrs borh while airborne and afrer rhey
are deposired on rhe ground. |or insrance, nirrous oxide
is longlived in rhe armosphere and conrribures ro rhe
humandriven enhancemenr of rhe greenhouse effecr rhar
likely warms rhe Earrh`s climare. Nirric oxide is an im
porranr precursor of acid rain and phorochemical smog.
Some of rhe human acriviries discussed above
affecr rhe armosphere direcrly. |or insrance, essenrially
all of rhe more rhan J0 Tg per year of fixed nirrogen
released in auromobile exhausrs and in orher emissions
from fossil fuel burning is emirred ro rhe armosphere as
nirric oxide. Orher acriviries indirecrly enhance emissions
ro rhe armosphere. |nrensive ferrilizarion of agriculrural
soils can increase rhe rares ar which nirrogen in rhe form
of ammonia is volarilized and losr ro rhe air. |r can also
speed rhe microbial breakdown of ammonium and nirrares
in rhe soil, enhancing rhe release of nirrous oxide. Even
in wild or unmanaged lands downwind of agriculrural or
indusrrial areas, rain or windborne deposirion of human
generared nirrogen can spur increased emissions of ni
rrogen gases from rhe soils.
N|trcas Ox|dc
Nirrous oxide is a very effecrive hearrrapping
gas in rhe armosphere, in parr because ir absorbs ourgo
ing radianr hear from rhe Earrh in infrared wavelengrhs
rhar are nor caprured by rhe orher major greenhouse
gases, warer vapor and carbon dioxide. by absorbing
and reradiaring rhis hear back roward rhe Earrh, nirrous
oxide conrribures a few percenr ro overall greenhouse
warming.
Alrhough nirrous oxide is unreacrive and long
lived in rhe lower armosphere, when ir rises inro rhe srraro
sphere ir can rrigger reacrions rhar deplere and rhin rhe
srrarospheric ozone layer rhar shields rhe Earrh from
damaging ulrravioler radiarion.
The concenrrarion of nirrous oxide in rhe armo
sphere is currenrly increasing ar rhe rare of rwo ro rhree
renrhs of a percenr per year. vhile rhar rise is clearly
documenred, rhe sources of rhe increase remain unre
solved. borh fossil fuel burning and rhe direcr impacrs
of agriculrural ferrilizarion have been considered and re
jecred as rhe major source. Rarher, rhere is a developing
consensus rhar a wide array of humandriven sources
conrribure sysremarically ro enrich rhe rerresrrial nirro
gen cycle. These dispersed sources" include ferrilizers,
nirrogenenriched groundwarer, nirrogensarurared for
esrs, foresr burning, land clearing, and even rhe manu
facrure of nylon, nirric acid, and orher indusrrial prod
ucrs.
The ner effecr is increased global concenrrarions
of a porenr greenhouse gas rhar also conrribures ro rhe
rhinning of rhe srrarospheric ozone layer.
N|tr|c Ox|dc aad Ammca|a
Unlike nirrous oxide, which is unreacrive in rhe
lower armosphere, borh nirric oxide and ammonia are
highly reacrive and rherefore much shorrer lived. Thus
6
Issues |n |co|ogy Numher 1 Spr|ng 1997
||||CTS ON TH| CAk8ON CYCL|
|ncreased emissions of airborne nirrogen have led
ro enhanced deposirion of nirrogen on land and in rhe
oceans. Thanks ro rhe ferrilizer effecrs of nirrogen in
srimularing planr growrh, rhis deposirion may be acring
ro influence rhe armosphere indirecrly by alrering rhe glo
bal carbon cycle.
Over much of rhe Earrh`s surface, rhe lushness
of planr growrh and rhe accumularion of sranding srocks
of planr marerial hisrorically have been limired by scanry
nirrogen supplies, parricularly in remperare and boreal
regions. Human acriviry has subsranrially increased rhe
deposirion of nirrogen over much of rhis area, which raises
imporranr quesrions: How much exrra planr growrh has
been caused by humangenerared nirrogen addirions' As
a resulr, how much exrra carbon has been srored in rer
resrrial ecosysrems rarher rhan conrriburing ro rhe rising
concenrrarions of carbon dioxide in rhe armosphere'
Answers ro rhese quesrions could help explain
rhe imbalance in rhe carbon cycle rhar has come ro be
known as rhe `missing sink.` The known emissions of
carbon dioxide from human acriviries such as fossil fuel
burning and deforesrarion exceed by more rhan 1,000
Tg rhe amounr of carbon dioxide known ro be accumu
laring in rhe armosphere each year. Could increased
growrh rares in rerresrrial vegerarion be rhe `sink` rhar
accounrs for rhe fare of much of rhar missing carbon'
Experimenrs in Europe and America indicare rhar
a large porrion of rhe exrra nirrogen rerained by foresr,
werland, and rundra ecosysrems srimulares carbon up
rake and srorage. On rhe orher hand, rhis nirrogen can
changes in rheir armospheric concenrrarions can be de
recred only ar local or regional scales.
Nirric oxide plays several crirical roles in armo
spheric chemisrry, including caralyzing rhe formarion of
phorochemical (or brownj smog. |n rhe presence of sun
lighr, nirric oxide and oxygen reacr wirh hydrocarbons
emirred by auromobile exhausrs ro form ozone, rhe mosr
dangerous componenr of smog. Croundlevel ozone has
serious derrimenral effecrs on human healrh as well as
rhe healrh and producriviry of crops and foresrs.
Nirric oxide, along wirh orher oxides of nirrogen
and sulfur, can be rransformed in rhe armosphere inro
nirric acid and sulfuric acid, which are rhe major compo
nenrs of acid rain.
Alrhough a number of sources conrribure ro ni
rric oxide emissions, combusrion is rhe dominanr one.
|ossil fuel burning emirs more rhan J0 Tg per year of
nirric oxide. Human burning of foresrs and orher planr
marerial may add abour 10 Tg, and global emissions of
nirric oxide from soils, a subsranrial fracrion of which are
humancaused, roral 5 ro J0 Tg per year. Overall, S0
percenr or more of nirric oxide emissions worldwide are
generared by human acriviries, and in many regions rhe
resulr is increased smog and acid rain.
|n conrrasr ro nirric oxide, ammonia acrs as rhe
primary acidneurralizing agenr in rhe armosphere, hav
ing an opposire influence on rhe acidiry of aerosols,
cloudwarer, and rainfall. Nearly 0 percenr of global
ammonia emissions are humancaused. Ammonia vola
rilized from ferrilized fields conrribures an esrimared 10
Tg per year: ammonia released from domesric animal
wasres abour JJ Tg: and foresr burning some 5 Tg.
7
||gure 4Human acriviries are responsible for a
large proporrion of rhe global emissions of ni
rrogenconraining rrace gases, including 40%
of rhe nirrous oxide, S0% or more of nirric ox
ide, and 0% of ammonia releases. The resulr is
increasing armospheric concenrrarions of rhe
greenhouse gas nirrous oxide, of rhe nirrogen
precursors of smog, and of biologically available
nirrogen rhar falls from rhe armosphere ro ferril
ize ecosysrems. Ammonia dara are from
Schlesinger and Harrley (199Jj, nirric oxide from
Delmas er al. (in pressj, and nirrous oxide from
Prarher er al. (1995j.
Issues |n |co|ogy Numher 1 Spr|ng 1997
Ammonia
(NH )
3
Nitric oxide
(NO)
Nitrous oxide
(N O)
2
20%
40%
60%
80%
100%
Human-Caused Global Nitrogen Emissions
rems. These impacrs firsr be
came apparenr in Europe almosr
rwo decades ago when scienrisrs
observed significanr increases in
nirrare concenrrarions in some
lakes and srreams and also ex
rensive yellowing and loss of
needles in spruce and orher co
nifer foresrs subjecred ro heavy
nirrogen deposirion. These ob
servarions led ro several field ex
perimenrs in rhe U.S. and Eu
rope rhar have revealed a com
plex cascade of effecrs ser in
morion by excess nirrogen in for
esr soils.
As ammonium builds up
in rhe soil, ir is increasingly con
verred ro nirrare by bacrerial
acrion, a process rhar releases
hydrogen ions and helps acidify
rhe soil. The buildup of nirrare
enhances emissions of nirrous
oxides from rhe soil and also en
courages leaching of highly wa
rersoluble nirrare inro srreams
or groundwarer. As rhese nega
rively charged nirrares seep
away, rhey carry wirh rhem posirively charged alkaline
minerals such as calcium, magnesium, and porassium.
Thus human modificarions ro rhe nirrogen cycle decrease
soil ferriliry by grearly acceleraring rhe loss of calcium
and orher nurrienrs rhar are viral for planr growrh. As
calcium is deplered and rhe soil acidified, aluminum ions
are mobilized, evenrually reaching roxic concenrrarions
rhar can damage rree roors or kill fish if rhe aluminum
washes inro srreams. Trees growing in soils replere wirh
nirrogen bur srarved of calcium, magnesium, and poras
sium can develop nurrienr imbalances in rheir roors and
leaves. This may reduce rheir phorosynrheric rare and
efficiency, srunr rheir growrh, and even increase rree
dearhs.
Nirrogen sarurarion is much furrher advanced
over exrensive areas of norrhern Europe rhan in Norrh
America because humangenerared nirrogen deposirion
is several rimes grearer rhere rhan in even rhe mosr ex
rremely affecred areas of Norrh America. |n rhe nirro
gensarurared ecosysrems of Europe, a subsranrial frac
rion of armospheric nirrare deposirs move from rhe land
inro srreams wirhour ever being raken up by organisms
also srimulare microbial decom
posirion and rhus releases of car
bon from soil organic marrer.
On balance, however, rhe car
bon uprake rhrough new planr
growrh appears ro exceed rhe
carbon losses, especially in for
esrs.
A number of groups
have arrempred ro calculare rhe
amounr of carbon rhar could be
srored in rerresrrial vegerarion
rhanks ro planr growrh spurred
by added nirrogen. The resulr
ing esrimares range from 100
ro 1,J00 Tg per year. The num
ber has rended ro increase in
more recenr analyses as rhe
magnirude of humandriven
changes in rhe nirrogen cycle
has become clearer. The mosr
recenr analysis of rhe global
carbon cycle by rhe |nrergovern
menral Panel on Climare Change
concluded rhar nirrogen depo
sirion could represenr a major
componenr of rhe missing car
bon sink.
More precise esrimares will become possible when
we have a more complere undersranding of rhe fracrion
of humangenerared nirrogen rhar acrually is rerained
wirhin various landbased ecosysrems.
NITkO6|N SATbkATION AND
|COSYST|M |bNCTIONIN6
There are limirs ro how much planr growrh can
be increased by nirrogen ferrilizarion. Ar some poinr,
when rhe narural nirrogen deficiencies in an ecosysrem
are fully relieved, planr growrh becomes limired by spar
siry of orher resources such as phosphorus, calcium, or
warer. vhen rhe vegerarion can no longer respond ro
furrher addirions of nirrogen, rhe ecosysrem reaches a
srare described as nirrogen sarurarion." |n rheory, when
an ecosysrem is fully nirrogensarurared and irs soils,
planrs, and microbes cannor use or rerain any more, all
new nirrogen deposirs will be dispersed ro srreams,
groundwarer, and rhe armosphere.
Nirrogen sarurarion has a number of damaging
consequences for rhe healrh and funcrioning of ecosys
8
||gure 5vild planrs living in narural ecosysrems,
such as rhis lupine, a nirrogenfixing planr, domi
nared rhe nirrogen cycle for millions of years. Hu
man producrion of nirrogen ferrilizer, burning of fossil
fuels, and inrensive culrivarion of legume crops now
adds as much nirrogen ro rerresrrial ecosysrems as
do all narural processes combined.
Issues |n |co|ogy Numher 1 Spr|ng 1997
rhis consrrainr. New supplies of nirrogen showered upon
rhese ecosysrems can cause a dramaric shifr in rhe domi
nanr species and also a marked reducrion in overall spe
cies diversiry as rhe few planr species adapred ro rake
full advanrage of high nirrogen our compere rheir neigh
bors. |n England, for example, nirrogen ferrilizers ap
plied ro experimenral grasslands have led ro increased
dominance by a few nirrogenresponsive grasses and loss
of many orher planr species. Ar rhe highesr ferrilizarion
rare, rhe number of planr species declined more rhan five
fold. |n Norrh America, similarly dramaric reducrions in
biodiversiry have been creared by ferrilizarion of grass
lands in Minnesora and California (|igures , S, and 9j.
|n formerly speciesrich hearhlands across vesrern Eu
rope, humandriven nirrogen deposirion has been blamed
for grear losses of biodiversiry in recenr decades.
|n rhe Nerherlands, high human popularion den
siry, inrensive livesrock operarions, and indusrries have
combined ro generare rhe highesr rares of nirrogen depo
sirion in rhe world. One welldocumenred consequence
has been rhe conversion of speciesrich hearhlands ro spe
ciespoor grasslands and foresr. Nor only rhe species
richness of rhe hearh bur also rhe biological diversiry of
rhe landscape has been reduced because rhe modified
planr communiries now resemble rhe composirion of com
muniries occupying more ferrile soils. The unique species
assemblage adapred ro sandy, nirrogenpoor soils is be
ing losr from rhe region.
or playing a role in rhe biological cycle.
|n conrrasr, in rhe norrheasrern U.S., increased
leaching of nirrares from rhe soil and large shifrs in rhe
nurrienr rarios in rree leaves generally have been observed
only in cerrain rypes of foresrs. These include highel
evarion sires rhar receive large nirrogen deposirs and sires
wirh shallow soils conraining few alkaline minerals ro
buffer acidificarion. Elsewhere in rhe U.S., rhe early srages
of nirrogen sarurarion have been seen in response ro el
evared nirrogen deposirion in rhe foresrs surrounding rhe
Los Angeles basin and in rhe |ronr Range of rhe Colo
rado Rockies.
Some foresrs have a very high capaciry ro rerain
added nirrogen, parricularly regrowing foresrs rhar have
been subjecred ro inrense or repeared harvesring, an ac
riviry rhar usually causes severe nirrogen losses. Over
all, rhe abiliry of a foresr ro rerain nirrogen depends on
irs porenrial for furrher growrh and rhe exrenr of irs cur
renr nirrogen srocks. Thus, rhe impacrs of nirrogen depo
sirion are righrly linked ro orher rapidly changing hu
mandriven variables such as shifrs in land use, climare,
and armospheric carbon dioxide and ozone levels.
||||CTS ON 8IODIv|kSITY AND TH| SP|CI|S MIX
Limired supplies of biologically available nirrogen
are a facr of life in mosr narural ecosysrems, and many
narive planr species are adapred ro funcrion besr under
9
||gure 6Deposirion of nirrogen from rhe armosphere is believed ro be responsible for rhe yellow
ing and loss of needles from conifers and for cases of foresr dieback, such as rhar shown here.
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Losses of biodiversiry driven by nirrogen deposi
rion can in rurn affecr orher ecological processes. Re
cenr experimenrs in Minnesora grasslands showed rhar
in ecosysrems made speciespoor by ferrilizarion, planr
producriviry was much less srable in rhe face of a major
droughr. Even in nondroughr years, rhe normal vagar
ies of climare produced much more yearroyear varia
rion in rhe producriviry of speciespoor grassland plors
rhan in more diverse plors.
||||CTS ON AObATIC
|COSYST|MS
H|stcr|ca| Chaaqcs |a Watcr
Chcm|strq
Nor surprisingly, ni
rrogen concenrrarions in sur
face warers have increased as
human acriviries have acceler
ared rhe rare of fixed nirrogen
being pur inro circularion. A
recenr srudy of rhe Norrh Ar
lanric Ocean basin by scien
risrs from a dozen narions es
rimares rhar movemenrs of ro
ral dissolved nirrogen inro
mosr of rhe remperarezone
rivers in rhe basin may have
increased by rwo ro J0fold
since preindusrrial rimes (|ig
ure 10j. |or rivers in rhe Norrh
Sea region, rhe nirrogen in
crease may have been six ro
J0fold. The nirrogen in
creases in rhese rivers are
highly correlared wirh human
generared inpurs of nirrogen ro rheir warersheds, and
rhese inpurs are dominared by ferrilizers and armospheric
deposirion.
|or decades, nirrare concenrrarions in many riv
ers and drinking warer supplies have been closely moni
rored in developed regions of rhe world, and analysis of
rhese dara confirms a hisroric rise in nirrogen levels in
rhe surface warers. |n 1,000 lakes in Norway, for ex
ample, nirrare levels doubled in less rhan a decade. |n
rhe Mississippi River, nirrares have more rhan doubled
since 1965. |n major rivers of rhe norrheasrern U.S.,
nirrare concenrrarions have risen rhree ro renfold since
rhe early 1900s, and rhe evidence suggesrs a similar
rrend in many European rivers.
Again nor surprisingly, nirrare concenrrarions in
rhe world`s large rivers rise along wirh rhe densiry of
human popularion in rhe warersheds. Amounrs of roral
dissolved nirrogen in rivers are also correlared wirh hu
man popularion densiry, bur roral nirrogen does nor in
crease as rapidly as rhe nirrare fracrion. Evidence indi
cares rhar wirh increasing human disrurbance, a higher
proporrion of rhe nirrogen in surface warers is composed
of nirrare.
|ncreased concenrrarions
of nirrare have also been ob
served in groundwarer in
many agriculrural regions, al
rhough rhe magnirude of rhe
rrend is difficulr ro derermine
in all bur a few wellcharac
rerized aquifers. Overall, rhe
addirions ro groundwarer
probably represenr only a
small fracrion of rhe increased
nirrare rransporred in surface
warers. However, groundwa
rer has a long residence rime
in many aquifers, meaning
rhar groundwarer qualiry is
likely ro conrinue ro decline as
long as human acriviries are
having subsranrial impacrs on
rhe nirrogen cycle.
High levels of nirrares in
drinking warer raise signifi
canr human healrh concerns,
especially for infanrs. Mi
crobes in an infanr`s sromach
may converr high levels of ni
rrare ro nirrire. vhen nirrire
is absorbed inro rhe bloodsrream, ir converrs oxygen
carrying hemoglobin inro an ineffecrive form called merh
emoglobin. Elevared merhemoglobin levels an anemic
condirion known as merhemoglobinemia can cause
brain damage or dearh. The condirion is rare in rhe U.S.,
bur rhe porenrial exisrs whenever nirrare levels exceed
U.S. Public Healrh Service srandards (10 milligrams per
lirerj.
N|trcqca aad Ac|d|[|cat|ca c[ lakcs
Nirric acid is playing an increasing role in rhe
acidificarion of lakes and srreams for rwo major reasons.
One is rhar mosr efforrs ro conrrol acid deposirion
which includes acid rain, snow, fog, misr, and dry depos
10
||gure 7Differenr rares of nirrogen addirion lead ro
marked changes in rhe planr and insecr species com
posirions and speicies diversiry of rhese plors of grass
land vegerarion in Minnesora. Each plor is 4m x 4m
(abour 1J fr x 1J frj, and has received experimenral
addirion of nirrogen (ammonium nirrarej since 19SJ.
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11
irs have focused on curring emissions of sulfur diox
ide ro limir rhe formarion of sulfuric acid in rhe armo
sphere. |n many areas, rhese efforrs have succeeded in
reducing inpurs of sulfuric acid ro soils and warer while
emissions of nirrogen oxides, rhe precursors of nirric acid,
have gone largely unchecked. The second reason is rhar
many warersheds in areas of moderare ro high nirrogen
deposirion appear ro be approaching nirrogen sarurarion,
and rhe increasingly acidified soils have lirrle capaciry ro
buffer acid rain before ir enrers srreams.
An addirional facror in many areas is rhar nirric
acid predominares among rhe polluranrs rhar accumu
lare in rhe winrer snowpack. Much of rhis nirric acid is
flushed our wirh rhe firsr barch of spring melrwarer, of
ren washing a sudden, concenrrared acid pulse" inro
vulnerable lakes.
Adding inorganic nirrogen ro freshwarer ecosys
rems rhar are also rich in phosphorus can eurrophy as
well as acidify rhe warers. borh eurrophicarion and acidi
ficarion generally lead ro decreased diversiry of borh planr
and animal species. |ish popularions, in parricular, have
been reduced or eliminared in many acidified lakes across
Scandinavia, Canada, and norrheasrern Unired Srares.
because rhe exrenr of nirrogensarurared ecosys
rems conrinues ro grow, along wirh humancaused nirro
gen deposirion, conrrols on sulfur dioxide emissions alone
clearly will nor be sufficienr ro decrease acid rain or pre
venr irs derrimenral effecrs on srreams and lakes. Euro
pean governmenrs already have recognized rhe impor
rance of nirrogen in acidifying soils and warers, and in
rergovernmenral efforrs are underway rhere ro reduce
emissions and deposirion of nirrogen on a regional basis.
Fatrcph|cat|ca |a Fstaar|cs aad Ccasta| Watcrs
One of rhe besr documenred and besr undersrood
consequences of human alrerarions of rhe nirrogen cycle
is rhe eurrophicarion of esruaries and coasral seas (|ig
ure 11 and 1Jj. |r is arguably rhe mosr serious human
rhrear ro rhe inregriry of coasral ecosysrems.
|n sharp conrrasr ro rhe majoriry of remperare
zone lakes, where phosphorus is rhe nurrienr rhar mosr
limirs primary producriviry by algae and orher aquaric
planrs and conrrols eurrophicarion, rhese processes are
conrrolled by nirrogen inpurs in mosr remperarezone es
ruaries and coasral warers. This is largely because rhe
narural flow of nirrogen inro rhese warers and rhe rare of
nirrogen fixarion by plankronic organisms are relarively
low while microbes in rhe sea floor sedimenrs acrively
release nirrogen back ro rhe armosphere.
vhen high nirrogen loading causes eurrophica
rion in srrarified warers where a sharp remperarure
gradienr prevenrs mixing of warm surface warers wirh
colder borrom warers rhe resulr can be anoxia (no
oxygenj or hypoxia (low oxygenj in borrom warers. borh
condirions appear ro be becoming more prevalenr in many
esruaries and coasral seas. There is good evidence rhar
since rhe 1950s or 1960s, anoxia has increased in rhe
balric Sea, rhe black Sea, and Chesapeake bay. Periods
||gure 8Narive grasslands in Minnesora ofren conrain J0
ro J0 or more planr species per square merer, as does rhis
plor. This plor is a conrrol" plor rhar received no nirrogen,
and rhar rerained irs original planr diversiry.
||gure 9Nirrogen addirion ro rhis plor, locared near rhar shown
in |igure S, led ro rhe loss of almosr all narive prairie species
and ro dominance by rhe weedy European quackgrass. |n
19SJ rhis plor looked much like rhe one shown in |igure S.
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year. There is some evidence rhar human alrerarion of
rhe nirrogen cycle could alrer biological processes in rhe
open ocean, bur rhere is no adequare frame of reference
againsr which ro evaluare any porenrial humandriven
change in marine nirrogen fixarion.
Chaaqcs |a l|m|t|aq kcscarccs
One consequence of humandriven changes in rhe
global nirrogen cycle is a shifr in rhe resources rhar limir
biological processes in many areas. Large amounrs of
nirrogen are now deposired on many ecosysrems rhar
were once nirrogen deficienr. The dominanr species in
rhese sysrems may have evolved wirh nirrogen limira
rion, and rhe ways rhey grow and funcrion and form
symbioric parrnerships may reflecr adaprarions ro rhis
limir. virh rhis limir removed, species musr operare un
der novel consrrainrs such as nowinadequare phospho
rus or warer supplies. How are rhe performance of or
ganisms and rhe operarion of larger ecological processes
affecred by changes in rheir chemical environmenr for
which rhey have no evolurionary background and ro which
rhey are nor adapred'
Capac|tq tc kcta|a N|trcqca
|oresrs and werlands vary subsranrially in rheir
capaciry ro rerain added nirrogen. |nreracring facrors
rhar are known ro affecr rhis capaciry include soil rex
rure, degree of chemical wearhering of soil, fire hisrory,
rare ar which planr marerial accumulares, and pasr hu
man land use. However, we srill lack a fundamenral un
dersranding of how and why nirrogenrerenrion processes
vary among ecosysrems much less how rhey have
changed and will change wirh rime.
of hypoxia have increased in Long |sland Sound, rhe Norrh
Sea, and rhe Karregar, resulring in significanr losses of
fish and shellfish.
Eurrophicarion is also linked ro losses of diver
siry, borh in rhe sea floor communiry including
seaweeds, seagrasses, and corals and among plank
ronic organisms. |n eurrophied warers, for example, nui
sance algae" may come ro dominare rhe phyroplankron
communiry. |ncreases in rroublesome or roxic algal blooms
have been observed in many esruaries and coasral seas
worldwide in recenr decades. During rhe 19S0s, roxic
blooms of dinoflagellares and brownride organisms
caused exrensive dieoffs of fish and shellfish in many
esruaries. Alrhough rhe causes are nor complerely un
dersrood, rhere is compelling evidence rhar nurrienr en
richmenr of coasral warers is ar leasr parrly ro blame for
such blooms.
MAJOk bNC|kTAINTI|S
Alrhough rhis reporr has focused on whar is
known abour humandriven changes ro rhe global nirro
gen cycle, major uncerrainries remain. Some of rhese
have been nored in earlier secrions. This secrion, how
ever, focuses on imporranr processes rhar remain so poorly
undersrood rhar ir is difficulr ro disringuish humancaused
impacrs or ro predicr rheir consequences.
Mar|ac N|trcqca F|xat|ca
Lirrle is known abour rhe unmodified nirrogen
cycle in rhe open ocean. Credible esrimares of nirrogen
fixarion by organisms in rhe sea vary more rhan renfold,
ranging from less rhan J0 ro more rhan J00 Tg per
1J
||gure 10Movemenrs of nirro
gen inro mosr of rhe remperare
zone rivers rhar empry inro rhe
Norrh Arlanric Ocean have in
creased by rwo ro J0fold since
preindusrrial rimes. Nirrogen in
creases in rhese rivers are highly
correlared wirh increasing human
generared nirrogen inpurs inro
rhe warersheds, parricularly fer
rilizer use and rising armospheric
deposirion of nirrogen. Modified
from Howarrh er al. (1996j.
Issues |n |co|ogy Numher 1 Spr|ng 1997
A|tcrat|ca c[ Dca|tr|[|cat|ca
|n large river basins, rhe majoriry of nirrogen rhar
arrives is probably broken down by denirrifying bacreria
and released ro rhe armosphere as nirrogen gas or ni
rrous oxide. Exacrly where mosr of rhis acriviry rakes
place is poorly undersrood, alrhough we know rhar srream
side areas and werlands are imporranr. Human acriviries
such as increased nirrare deposirion, dam building, and
rice culrivarion have probably enhanced denirrificarion,
while draining of werlands and alrerarion of riparian eco
sysrems has probably decreased ir. bur rhe ner effecr of
human influence remains uncerrain.
Natara| N|trcqca Cqc||aq
|nformarion on rhe rare of nirrogen depo
sirion and loss in various regions prior ro exrensive
human alrerarions of rhe nirrogen cycle remains
parchy. |n parr, rhis reflecrs rhe facr rhar all of rhe
Earrh already is affecred ro some degree by hu
man acriviry. Neverrheless, srudies in remore re
gions of rhe Sourhern Hemisphere illusrrare rhar
rhere is srill valuable informarion ro be garhered on
areas rhar have been minimally alrered by humans.
|bTbk| PkOSP|CTS AND
MANA6|M|NT OPTIONS
Fcrt|||zcr Usc
The grearesr humandriven increases in glo
bal nirrogen supplies are linked ro acriviries inrended
ro boosr food producrion. Modern inrensive agri
1J
culrure requires large quanriries of nirrogen ferril
izer: humaniry, in rurn, requires inrensive agricul
rure ro supporr a growing popularion rhar is pro
jecred ro double by rhe end of rhe nexr cenrury.
Consequenrly, rhe producrion and applicarion of
nirrogen ferrilizer has grown exponenrially, and rhe
highesr rares of applicarion are now found in some
developing counrries wirh rhe highesr rares of popu
larion growrh. One srudy predicrs rhar by rhe
year J0J0, global producrion of nirrogen ferril
izer will increase from a currenr level of abour S0
Tg ro 1J4 Tg per year.
Currailing rhis growrh in nirrogen ferrilizer pro
ducrion will be a difficulr challenge. Neverrheless,
rhere are ways ro slow rhe growrh of ferrilizer use
and also ro reduce rhe mobiliry and hence rhe
regional and global impacrs of rhe nirrogen
rhar is applied ro fields.
One way ro reduce rhe amounr of ferrilizer used
is ro increase irs efficiency. Ofren ar leasr half of rhe
ferrilizer applied ro fields is losr ro rhe air or warer. This
leakage represenrs an expensive wasre ro rhe farmer as
well as a significanr driver of environmenral change. A
number of managemenr pracrices have been idenrified
rhar can reduce rhe amounrs of ferrilizer used and cur
losses of nirrogen ro rhe air and warer wirhour sacrific
ing yields or profirs (and in some cases, increasing rhemj.
|or insrance, one commercial sugar cane planrarion in
Hawaii was able ro cur nirrogen ferrilizer use by one rhird
and reduce losses of nirrous oxide and nirric oxide ren
||gure 11The borromdwelling planrs of a marine ecosysrem
rhar received narural rares of nirrogen addirion. Nore rhe high
diversiry of rhese planrs and rheir spacing.
|igure 1JThe borromdwelling planrs of a marine ecosysrem rhar
received high rares of nirrogen inpur. Nore rhar rhere are few
planr species, and rhar rhe leaves of rhese are covered wirh a
rhick layer of algae.
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Issues |n |co|ogy Numher 1 Spr|ng 1997
ing world. One srudy predicrs rhar producrion of nirro
gen oxides from fossil fuels will more rhan double in rhe
nexr J5 years, from abour J0 Tg per year ro 46 Tg.
Reducing rhese emissions will require improvemenrs in
rhe efficiency of fuel combusrion as well as in rhe inrer
ceprion of airborne byproducrs of combusrion. As wirh
improvemenrs in ferrilizer efficiency, ir will be parricularly
imporranr ro rransfer efficienr combusrion rechnologies
ro developing counrries as rheir economies and indus
rries grow.
CONCLbSIONS
Human acriviries during rhe pasr cenrury have
doubled rhe narural annual rare ar which fixed nirrogen
enrers rhe landbased nirrogen cycle, and rhe pace is likely
ro accelerare. Serious environmenral consequences are
already apparenr. |n rhe armosphere, concenrrarions of
rhe greenhouse gas nirrous oxide and of rhe nirrogen
precursors of smog and acid rain are increasing. Soils in
many regions are being acidified and srripped of nurri
enrs essenrial for conrinued ferriliry. The warers of srreams
and lakes in rhese regions are also being acidified, and
excess nirrogen is being rransporred by rivers inro esru
aries and coasral warers. |r is quire likely rhar rhis un
precedenred nirrogen loading has already conrribured ro
fold by dissolving rhe ferrilizer in irrigarion warer, deliv
ering ir below rhe soil surface, and riming mulriple appli
carions ro meer rhe needs of rhe growing crop. This
knowledgeinrensive sysrem also proved more profirable
rhan broadcasring fewer, larger applicarions of ferrilizer
onro rhe soil surface. The widespread implemenrarion of
such pracrices, parricularly in developing regions, should
be a high prioriry for agronomisrs as well as ecologisrs
since improved pracrices provide an opporruniry ro re
duce rhe cosrs of food producrion while slowing rhe rare
of global change.
There are also ways ro prevenr rhe nirrogen rhar
leaches from ferrilized farmland from reaching srreams,
esruaries and coasral warers where ir conrribures ro
eurrophicarion. |n many regions, agriculrural lands have
been expanded by channelizing srreams, clearing ripar
ian foresrs, and draining werlands. Yer rhese areas serve
as imporranr narural nirrogen rraps. Resrorarion of wer
lands and riparian areas and even consrrucrion of arrifi
cial werlands have been shown ro be effecrive in prevenr
ing excess nirrogen from enrering warers.
Fcss|| Fac| 8ara|aq
The second major source of humanfixed nirro
gen is fossil fuel burning. |r, roo, will increase markedly
as we enrer rhe nexr cenrury, parricularly in rhe develop
14
||gure 13Human acriviries, such as ferrilizer producrion, growing legume crops, and burning of fossil fuels, are now
of equal or grearer magnirude rhan narural processes in rhe nirrogen cycle. Human dominarion of rhe nirrogen cycle
impacrs rhe funcrioning of many rerresrrial and aquaric ecosysrems, including seemingly prisrine habirars such as rhis
alpine ecosysrem.
Issues |n |co|ogy Numher 1 Spr|ng 1997
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limirarion on land and in rhe sea: How can ir occur' bio
geochemisrry 1J:S115.
Abcat thc Paac| c[ Sc|cat|sts
This reporr presenrs rhe consensus reached by a
panel of eighr scienrisrs chosen ro include a broad array
of experrise in rhis area. This reporr underwenr peer
review and was approved by rhe board of Edirors of |s
sues in Ecology. The affiliarions of rhe members of rhe
panel of scienrisrs are:
Dr. Perer M. virousek, Panel Chair, Deparrmenr of bio
logical Sciences, Sranford Universiry, Sranford, CA 94J05
longrerm declines in coasral fisheries and accelerared
losses of planr and animal diversiry in borh aquaric and
landbased ecosysrems. |r is urgenr rhar narional and
inrernarional policies address rhe nirrogen issue, slow rhe
pace of rhis global change, and moderare irs impacrs.
|Ok |bkTH|k IN|OkMATION
This reporr summarizes rhe findings of our panel.
Our full reporr, which is being published in rhe journal
Ecc|cq|ca| ^pp||cat|cas (volume , Augusr 199j, dis
cusses and cires more rhan 140 references ro rhe pri
mary scienrific lirerarure on rhis subjecr. |rom rhar lisr
we have chosen rhose below as illusrrarive of rhe scien
rific publicarions and summaries upon which our reporr
is based.
Aber, 1.D. 199J. Nirrogen cycling and nirrogen sarura
rion in remperare foresr ecosysrems. Trends in Ecology
and Evolurion :JJ0JJJ.
berendse, |., R. Aerrs, and R. bobbink. 199J. Armo
spheric nirrogen deposirion and irs impacr on rerresrrial
ecosysrems. Pp. 1041J1 in C.C. vos and P. Opdam
(edsj, Landscape Ecology of a Srressed Environmenr.
Chapman and Hall, England.
Cole, 1. 1., b. L. Peierls, N. |. Caraco, and M. L. Pace.
199J. Nirrogen loadings of rivers as a humandriven pro
cess. Pages 14115 in M. 1. McDonnell and S. T. A.
Picker (eds.j, Humans as Componenrs of Ecosysrems:
The Ecology of Subrle Human Effecrs and Populared Ar
eas. Springerverlag, NY.
DOE (Deparrmenr of Environmenr, UKj. 1994. |mpacrs
of Nirrogen Deposirion in Terresrrial Ecosysrems. Techni
cal Policy branch, Air Oualiry Div., London.
Calloway, 1. N., v. H. Schlesinger, H. Levy ||, A. Michaels,
and 1. L. Schnoor. 1995. Nirrogen fixarion: armospheric
enhancemenrenvironmenral response. Clobal bio
geochemical Cycles 9:JJ5J5J.
Howarrh, R. v., C. billen, D. Swaney, A. Townsend, N.
1aworski, K. Lajrha, 1. A. Downing, R. Elmgren, N.
Caraco, T. 1ordan, |. berendse, 1. |reney, v. Kudeyarov,
P. Murdoch, and Zhu Zhaoliang. 1996. Regional nirro
gen budgers and riverine N 6 P fluxes for rhe drainages
ro rhe Norrh Arlanric Ocean: Narural and human influ
ences. biogeochemisrry J5: 51J9.
15
Issues |n |co|ogy Numher 1 Spr|ng 1997
ogy, Evolurion and behavior, Universiry of Minnesora,
Sr. Paul, MN 5510S609. Email: rilman@lrer.umn.edu
8oarJ memhers
Dr. Srephen Carpenrer, Cenrer for Limnology, Universiry
of visconsin, Madison, v| 5J06
Dr. Deborah 1ensen, The Narure Conservancy, 1S15
Norrh Lynn Srreer, Arlingron, vA JJJ09
Dr. Simon Levin, Deparrmenr of Ecology 6 Evolurionary
biology, Princeron Universiry, Princeron, N1
0S544100J
Dr. 1ane Lubchenco, Deparrmenr of Zoology, Oregon
Srare Universiry, Corvallis, OR 9JJ1J914
Dr. 1udy L. Meyer, |nsrirure of Ecology, The Universiry of
Ceorgia, Arhens, CA J060JJJ0J
Dr. Cordon Orians, Deparrmenr of Zoology, Universiry
of vashingron, Searrle, vA 9S195
Dr. Lou Pirelka, Appalachian Environmenral Laborarory,
Cunrer Hall, |rosrburg, MD J15JJ
Dr. villiam Schlesinger, Deparrmenrs of borany and
Ceology, Duke Universiry, Durham, NC J0S
0J40
Add|t|caa| Ccp|cs
To receive addirional copies of rhis reporr, please
conracr:
Public Affairs Office
Ecological Sociery of America
J010 Massachuserrs Avenue, Nv
Suire 400
vashingron, DC J00J6
esahq@esa.org
(J0Jj SJJSJ
This version of rhe reporr is also available elecrronically
ar hrrp://www.sdsc.edu/-ESA/.
Also available for a small fee are reprinrs of our
full reporr, published in rhe journal Ecc|cq|ca| ^pp||ca
t|cas (volume , Augusr 199j wirh derailed cirarions ro
rhe original scienrific lirerarure. Conracr rhe Ecological
Sociery of America ar rhe above lisred address for more
informarion.
Special rhanks ro rhe U.S. Environmenral Prorec
rion Agency for supporring prinring and disrriburion of
rhis documenr.
Dr. 1ohn Aber, Complex Sysrems Research Cenrer, |nsri
rure for rhe Srudy of Earrh, Oceans and Space, Univer
siry of New Hampshire, Durham, NH 0JSJ4J5J5
Dr. Roberr v. Howarrh, Secrion of Ecology and Sysrem
arics, Corson Hall, Cornell Universiry, |rhaca, NY 14S5J
Dr. Cene E. Likens, |nsrirure of Ecosysrem Srudies, Cary
Arborerum, Millbrook, NY 1J545
Dr. Pamela A. Marson, Soil Science, Universiry of Cali
fornia, berkeley, berkeley, CA 94J0
Dr. David v. Schindler, Deparrmenr of biological Sciences,
Universiry of Alberra, Edmonron, Alberra, T6C JE9,
CANADA
Dr. villiam H. Schlesinger, Deparrmenrs of borany and
Ceology, Duke Universiry, Durham, NC J0S0J40
Dr. David Tilman, Deparrmenr of Ecology, Evolurion and
behavior, Universiry of Minnesora, Sr. Paul, MN 5510S
609
Abcat thc Sc|cacc Wr|tcr
Yvonne baskin, a science wrirer, edired rhe re
porr of rhe panel of scienrisrs ro allow ir ro more effec
rively communicare irs findings wirh nonscienrisrs.
Abcat Issacs |a Fcc|cqq
|ssaes |a Ecc|cqq is designed ro reporr, in lan
guage undersrandable by nonscienrisrs, rhe consensus
of a panel of scienrific experrs on issues relevanr ro rhe
environmenr. |ssues in Ecology is supporred by a Pew
Scholars in Conservarion biology granr ro David Tilman
and by rhe Ecological Sociery of America. All reporrs
undergo peer review and musr be approved by rhe ediro
rial board before publicarion.
Acknow|eJgements
This series was inspired by Dr. Ron Pulliam, who
firsr proposed rhe idea, and Dr. 1udy Meyer who chaired
an Ecological Sociery of America commirree rhar devel
oped rhe conceprs and convinced ESA ro pursue ir. ve
grarefully acknowledge rheir conrriburions. ve also rhank
|airh Kearns for her assisrance wirh design and produc
rion of rhis issue.
|J|tor|a| 8oarJ ot Issacs |a Fcc|cqq
Dr. David Tilman, EdirorinChief, Deparrmenr of Ecol
16
Issues |n |co|ogy Numher 1 Spr|ng 1997