Table Of ContentDevelopments in Plant and Soil Sciences 
Volume 6 
Also in  this series 
1.  J. Monteith and C. Webb, eds., 
Soil Water and Nitrogen in Mediterranean type Environments. 
1981. ISBN 90·247·2406·6 
2.  J.C. Brogan, ed., 
Nitrogen Losses and Surface Run·off from Landspreading of Manures. 
1981. ISBN 90·247·2471·6 
3.  J.D. Bewley, ed., 
Nitrogen and Carbon Metabolism. 
1981. ISBN 90·247·24724 
4.  R.  Brouwer, I. Ga~parfkova, J. Kolek and B.C.  Loughman, 
Structure and Function of Plant Roots. 
1981. ISBN 90·247·2405·8 
5.  Y.R. Dommergues and H.G. Diem, eds., 
Microbiology of Tropical Soils and Plant Productivity. 
1982. ISBN 90·247·2624·7 
In preparation 
J.R.  Freney and J.R. Simpson, eds., 
Gaseous Losses of Nitrogen from  Plant·Soil Systems. 
Series ISBN 90·247·2405·8
NITROGEN CYCLING IN ECOSYSTEMS OF LATIN AMERICA 
AND THE CARIBBEAN
United Nations Environment 
Program  (UNEP) 
Nitrogen Cycling in Ecosystems of 
Latin America and the  Caribbean 
edited by 
G.P.  ROBERTSON 
R.HERRERA 
T.  ROSSWALL 
Reprinted  from  Plant and  Soil  Vol.  67  (1982) 
1982 
MARTINUS NIJHOFF / DR W.  JUNK PUBLISHERS 
THE  HAGUE / BOSTON / LONDON
Proceedings of a  regional  workshop arranged by the SCOPE / UNEP International Nitrogen Unit 
of the Royal Swedish Academy of Sciences under UNEP contract FP /1303·78·01  (1330) at the 
Centro Internacional de Agricultura Tropical (CIAT), Cali, Colombia, 16·21  March,  1981; 
a meeting sponsored by  SCOPE, UNEP, MAB and COSTED. 
Distributors 
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Kluwer Boston, Inc. 
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Hingham, MA 02043 
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for all other countries 
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Distribution Center 
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l.ibrary  of Congress  Cataloging  in  Publication  Data 
Main  entry  under  title: 
Nitrogen  cycling  in Latin  American  and  Caribbean 
escosystems. 
(Developments  in  plant  and  soil  sciences  ;  v.  6) 
"Reprinted  from  Plant  and  soil,  vol.  67." 
Papers  from  a  workshop  held  at  Ciat,  Cali,  Colombia 
in March  1981,  which  was  arranged  by  the  SCOPE/UNEP  , 
International Nitrogen  Unit. 
Includes  bibliographies  and  index. 
1.  Nitrogen  cycle--Latin America--Congresses. 
2.  Nitrogen  cycle--Caribbean  area--Congresses. 
3.  Agricultural  ecology--Latin  America--Congresses. 
4.  Agricultural  ecology--Caribbean  area--Congresses. 
I.  Robertson,  G.  P.  II.  Herrera,  R.  III.  Rosswall,  T. 
(Thomas)  IV.  SCOPE/UNEP  International Nitrogen  Unit. 
V.  Series. 
QJU06.5.N57  1982  631.4'17  82-12631 
ISBN  90-247-2719-7 
ISBN-13:  978-94-009-7641-2  e-ISBN-13: 978-94-009-7639-9 
DOl:  10.1007/978-94-009-7639-9 
Copyright  ©  1982 by Martinus Nijhoff / Dr W.  Junk Publishers, The Hague. 
Softcover reprint of the hardcover 1st edition 1982 
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, 
or transmitted in any form or by any means, mechanical, photocopying, recording, or 
otherwise, without the prior written permission of the publishers, 
Martinus Nijhoff / Dr W.  Junk Publishers, P.O.Box 566, 2501 CN The Hague, The Netherlands.
Preface 
The large and rapidly expanding body ofliterature related to nitrogen cycling 
in  both  managed  and  native  terrestrial  ecosystems  reflects  the  importance 
accorded to the behaviour of this vital and often limiting nutrient. Research at the 
organism,  ecosystem  and  landscape  levels  commonly  addresses  questions 
concerning nitrogen acquisition,  internal cycling and  retention.  Goals for  this 
research include increased agricultural productivity and a better understanding 
of human impact on local,  regional and global nitrogen cycles. 
Nitrogen  cycle  research  in  tropical  regions  has  a  long  and  distinguished 
history.  Research  on different aspects of nitrogen cycling  in  ecosystems  of the 
tropics has been carried out in many regions. In relatively few instances has there, 
however, been a focus on the biogeochemical cycles at the ecosystem level. The 
meeting  resulting  in  this  volume  was  an  attempt  to  bring  together  existing 
information  on  nitrogen  cycling  in  ecosystems  of  Latin  America  and  the 
Caribbean and discuss this in an ecosystem context. 
The papers represent the proceedings of a workshop on Nitrogen Cycling in 
Ecosystems of Latin America and the Caribbean, the third workshop on nitrogen 
cycling within particular regions organized by the SCOPEjUNEP International 
Nitrogen  Unit  of  the  Royal  Swedish  Academy  of Sciences,  Stockholm.  The 
purpose of the  workshop was  fivefold:  1)  to  emphasize  the  importance of the 
nitrogen cycle in the different ecosystems of the region, 2) to provide a forum for 
scientists  from  the  region  to  present  papers describing ongoing nitrogen-cycle 
research,  3)  to  compile  available  data  into  coherent  nitrogen  budgets  for  the 
region's main ecosystems, and 5)  to define nitrogen-cycle research priorities for 
the region. Previous workshops have been held in West Africa 1 and in Southeast 
Asia 2.  The  three  workshops  have  been  supported  by  UNEP  under  contract 
FPj1303-78.01(1330). 
The  present  workshop  was  held  16-21  March,  1981,  at  CIAT  (Centro 
Internacional  de  Agricultura  Tropical)  in  Cali,  Colombia.  Three  days  of 
symposia  and  contributed  paper  sessions  were  followed  by  two  days  of 
workgroup discussions organized around major ecosystems of the region. These 
included shifting cultivation and traditional agroecosystems, sugarcane, cereal 
and grain crops, coffee and cacao plantations, savannas and shrublands, forests, 
and  wetlands  and  aquatic  systems.  Workgroups  were  charged  with  building 
informal nitrogen budgets of the respective systems and thereby summarizing the 
current state of knowledge regarding nitrogen cycles in each system. They were 
also asked to discuss research priorities, which were later reviewed by the plenary 
session. These priority rankings will,  we  hope, be useful  for  efficiently focusing 
increasingly  scarce  research  resources  on  important  but  little-understood 
nitrogen-cycle processes. 
v
The volume contains most ofthe papers presented at the meeting and the work 
group reports. Three additional papers by scientists from  the region unable to 
participate in the meeting are also included. A number of papers were originally 
presented in Spanish or Portuguese. In order to ensure as  large an audience as 
possible  for  these  reports  we  decided  to  publish  all  papers  in  English  with  a 
Spanish summary. 
Co-sponsors of the meeting apart from SCOPE and UNEP included the Man 
and the Biosphere (MAB) programme of Unesco, and the Committee on Science 
and Technology in Developing Countries (COSTED). We are greatly indebted to 
all  the sponsoring organizations for  their interest and support. The organizers 
also  extend particular  thanks  to CIA T  Director General J.  L. Nickel  and  his 
hospitable staff, and also to the simultaneous translators. 
We  are  also  indebted  to  Britta  Myrvik,  Gudrun  Sunnerstrand  and  Peter 
Wigren for artwork revisions and to Dina Soderstrom and Gun Martinsson for 
typing the final manuscript. 
Despite minor difficulties  with communicating in  three  languages, we  think 
most participants will agree that the workshop was a success and that its major 
objectives were well-met. 
East Lansing and Uppsala, April,  1982 
G.  P. Robertson, R.  Herrera and T.  Rosswall 
References 
Rosswall  T  (Ed.)  1980  Nitrogen  Cycling  in  West  African  Ecosystems.  Stockholm:  Royal 
Swedish Academy of Sciences, 450 p. 
2  Wetselaar R,  Simpson J Rand Rosswall T (Eds.)  1981  Nitrogen Cycling in South-East Asian 
Wet Monsoonal Ecosystems. Canberra: Australian Academy of Sciences. 216 p. 
VI
Contents 
Preface  V 
A. A. Franco, and D.  N.  Munns,  Plant assimilation and nitrogen cycling  I 
T.  Rosswall, Microbiological regulation of the biogeochemical nitrogen cycle  15 
P.  S. C.  Rao, R. E.  Jessup and A. G. Hornsby, Simulation of nitrogen in agro-ecosystems: Criteria for model selection and use  35 
D.  J.  Greenwood, Nitrogen supply and crop yield: The global scene  45 
E.  Sanhueza, The role of the atmosphere in nitrogen cycling  61 
G.  P.  Robertson, Regional nitrogen budgets: Approaches and problems  73 
A. J.  Holding,  Some priority research areas in nitrogen studies  81 
P.  A. Simchez, Nitrogen in shifting cultivation systems of Latin America  91 
S. R. Gliessman, Nitrogen distribution in several traditional agro-ecosystems in the humid tropical lowlands of south-eastern 
Mexico  105 
J.  D. H. Lambert and J.  T.  Amason, Nitrogen distribution in hybrid and local corn varieties and its possible relationship to a 
declining soil nitrogen pool under shifting agriculture at Indian Church, Belize  119 
R. L.  Victoria, P. L. Libardi, K. Reichardt and E. Matsui, 15N-urea transport and transformation in two deforested Amazonian 
soils under laboraiory conditions  129 
A. P.  Ruschel and P.  B.  Vose,  Nitrogen cycling in sugarcane  139 
S.  Valdivia  Vega,  Nitrogen gains and losses in  sugarcane (Saccharum sp.) agro-ecosystems on the coast of Peru  147 
M. N.  Versteeg, I.  Zipori, J.  Medina and H.  Valdivia, Potential growth of alfalfa (Medicago sativa L.) in the desert of Southern 
Peru and its  response to high NPK fertilization  157 
N.  Ahmad, E.  D.  Reid, M. Nkrumah, S. M. Griffith and L. Gabriel, Crop utilization and fixation of added ammonium in soils of 
the West Indies  167 
M.  A.  Lazzari, Distribution of l'N fertilizer  in field-Iysimeters  sown with garlic (Allium sativum) and foxtail millet  (Setaria 
italica)  187 
P.  L.  Libardi, R.  L.  Victoria,  K.  Reichardt and A.  Cervellini, Nitrogen cycling in a  l'N-fertilized bean (Phaseolus vulgaris L.) 
crop  193 
K.  R.  Reddy, Nitrogen cycling in a flooded-soil ecosystem planted to rice (Oryza sativa L.)  209 
E.  Schalscha B. and I.  Vergara F.,  The nitrogen balance of vegetable crops irrigated with untreated effluent  221 
J.  R. Jardim Freire,  Research into the Rhizobium/Leguminosae symbiosis in Latin America  227 
E.  Bomemisza, Nitrogen cycling in coffee plantations  241 
J.  Aranguren, G.  Escalante and R. Herrera, Nitrogen cycle of tropical perennial crops under shade trees:  I. Coffee  247 
J.  Aranguren, G.  Escalante and R.  Herrera,  Nitrogen cycle of tropical perennial crops under shane trees:  II. Cacao  259 
M.  B.  M. Santana and P.  Cabala-Rosand, Dynamics of nitrogen in a shaded cacao plantation  271 
J.  P.  Roskoski, Nitrogen fixation  in a Mexican coffee plantation  283 
J.  Pereira,  Nitrogen cycling in South American savannas  293 
E.  Medina,  Nitrogen balance in the Trachypogon grasslands of Central Venezuela  305 
R. H. Braun Wilke, Net primary productivity and nitrogen and carbon distribution in two xerophytic communities of central-
west Argentina  315 
C.  Jordan,  W.  Caskey,  G.  Escalante,  R.  Herrera,  F.  Montagnini,  R.  Todd and C.  Uhl,  The nitrogen cycle in a  'Terra Firme' 
rainforest on oxisol in  the Amazon territory of Venezuela  325 
J.  T.  Amason and J.  D.  H.  Lambert, Nitrogen cycling in  the seasonally dry forest zone of Belize. Central America  333 
P.  W.  Runde!, E.  T.  Nilsen,  M.  R.  Shariji,  R.  A.  Virginia,  W.  M. Jarrell,  D. H.  Kohl and G.  B. Shearer, Seasonal dynamics of 
nitrogen cycling for a Prosopis woodland in  the Sonoran Desert  343 
U.  Irmler,  Litterfall and nitrogen turnover in an Amazonian blackwater inundation forest  355 
R. E.  Cisternas and L. R.  Yates, Nitrogen in litterfall and precipitation and its release during litter decomposition in the Chilean 
piedmont matorral  359 
E.  Salali,  R.  Sylvester-Bradley and R. L.  Victoria,  Regional gains and losses of nitrogen  in  the Amazon basin  367 
S.  Cabrera S.  and V.  Montrcino B .. Eutrophy in Lake Aculeo. Chile  377 
Work group reports 
Shifting cultivation and traditional agriculture  389 
Sugarcane  395 
Cereal and grain crops  399 
Coffee and cacao plantations  403 
Savannas and shrublands  409 
Latin American forests  415 
Wetlands (including rice) and aquatic systems  421 
List of participants  425 
Index  429 
VII
Plant and Soil 67,  1-13 (1982).  0032-079X/82/0671-0001$01.95.  SU-01 
© 1982 Martinus Nijhoff/Dr W Junk Publishers,  The Hague. 
Plant assimilation and nitrogen cycling 
Asimilacion de nitrogeno por  las plantas y el cicio de este elemento 
A.A.FRANCO 
Empresa Brasileira de  Pesquisa  Agropecuaria (EM BRAP A), SN LCS-P F BN -km 47, Seropedica,  Rio 
de  Janeiro 23460, Brazil 
and D. N. MUNNS 
Department of Land, Air and Water Resources, University of California, Davis, California 95616, USA 
Key· words  N-assimilation  N-cycling  N-fertilization  N 2-fixation  Nitrogenase  N-
mineralization  N-reductase  pH changes 
Abstract  Nitrogen, an abundant and yet limiting nutrient for crop and food production, enters the 
plant as nitrate or ammonium, or as dinitrogen through biological fixation by procaryotes associated 
with  the plant. Nitrogen incorporation into the soil-plant-animal system is ultimately restricted by 
rates of biological and industrial fixation.  Biological fixation conserves fossil fuel, but fertilization is 
preferred in most present agriculture. Nitrogen-metabolism research has the practical objectives of 
allowing more efficient N-fertilizer utilization by plants, including those that fix N 2  but benefit from 
fertilizer-N supplements. 
Nitrogen accumulation by harvested crops results in changes in  soil acidity, with the direction of 
change  depending  on  the  N-source.  There  is  little  information  on  long-term  effects  of crop  N-
nutrition  on  acidity,  and acidity  is  a  critical  factor  that  affects  agricultural  productivity  in  many 
tropical soils.  Thus,  plant control of pH  and  the acid/base balance in  the soil  as a  consequence of 
nitrogen uptake and assimilation are important areas of future research. 
Resumen  EI nitr6geno, abundante pero sin embargo limit ante para los cultivos, entra en las plantas 
en  forma  de  nitrato  0  amonio  0  es  incorporado  al  sistema  a  traves  de  fijaci6n  biol6gica.  La 
incorporaci6n del  nitr6geno  al  sistema suelo-planta-animal est a limitado  por las  tasas de  fijaci6n 
biol6gica e industrial. La primera ahorra energia fosil pero la segunda fuente es la predominante en la 
agricultura moderna. La investigaci6n del metabolismo de nitr6geno tiene objetivos practicos tales 
como el  permitir un uso mas eficiente de los  fertilizantes nitrogenados por los cultivos, incluyendo 
aquellos que puedan fijar N 2  pero se benefician de suplementos de fertilizantes nitrogenados. 
La acumulaci6n de nitrogeno en los cultivos trae como consecuencia cam bios de acidez en el suelo 
cuya direccion depende de la forma de nitrogeno utilizada. Aun existe poca informacion sobre los 
efectos  a  largo  plazo  de  la  fertilizacion  nitrogenada  sobre  la  acidez  del  suelo,  factor  que  es 
determinante de la productividad de muchos suelos en los tropicos. Asi pues, el control de pH por las 
plantas y el  balance de acidez en el  suelo son areas de interes para futuras  investigaciones. 
Introduction 
Nitrogen is abundant but is  the nutrient that most often limits crop and food 
production.  A  crop  can  accumulate  up  to  800 kg  N  ha - 1  yr - \  most  non-
fertilized  tropical  soils  under agriculture deliver  less  than  50 kg  N  ha - 1  yr - 1 
(Sanchez 53 ).
2  FRANCO AND MUNNS 
The  availability  of  nitrogen  in  soil  is  limited  by  rates  of  organic  matter 
decomposition  unless  availability  is  enhanced  by  biological  N rfixation  or 
additions of fertilizer-N. Optimizing these  three sources for crop production is 
desirable for economic and ecological reasons. 
In this paper we discuss the turnover and availability of nitrogen in the soil and 
plant uptake and nitrogen assimilation, including the special case oflegumes and 
the  balance  of pH  as  a  consequence  of N  uptake.  pH  balance  is  of special 
importance because of its impact on the already acid soils found in most of the 
tropics. 
Nitrogen in the soil 
Forms of nitrogen 
Inorganic  nitrogen  in  soil  has  three  main  sources:  soil  organic  matter, 
atmospheric N z, and N-fertilizer.  During decomposition  of organic matter in 
most agricultural  soils, excess NH4 +  not utilized by microbes is  released,  and 
subsequently usually oxidized mostly by autotrophic bacteria to NOz - and then 
N03 -. Nitrite does  not  usually  accumulate except  temporarily  in  the  special 
situation  where  pH  is  above  7  and  excess  NH40H (or  urea  hydrolysing  to 
NH40H) together inhibit the NO z - oxidizers. 
When atmospheric N z is fixed, the first form of combined nitrogen to appear is 
NH4 + , and most of it is immediately assimilated into organic forms so that very 
little is exuded to soil. Both organic and exuded NH4 + and fertilizer-N follow the 
same path as N derived from organic matter. 
Total  nitrogen  in  soil  varies  with  soil  organic  matter content;  soil  organic 
matter usually contains ca.  5%  N.  In surface mineral  soils,  values  of 0.03%  N 
(Vertisol  from  Sudan)  to 0.69%  N  (Oxisol  from  Brazil) have  been  reported 53. 
Levels  of nitrate  to 0.006%  N03-N (60mg N03-Nkg dry  soil- 1)  have  been 
noted  by  Chapman 15 ,  while  ammonium  is  usually  much  less  except  in 
waterlogged soils after the addition ofNH4 + -N, urea, or nitrification inhibitors. 
Factors affecting nitrogen release 
In the absence offertilizer N, the major source of fixed nitrogen the soil supplies 
to the plant is  from  soil organic matter undergoing decomposition. 
As  illustrated in Fig.  1,  the availability of nitrogen to plants depends on the 
amount and  type  of organic matter present, and on the presence of microbial 
populations and conditions favoring their activity. In general, microbial activity 
is  favored  under  those  conditions  that  are  optimal  for  plant  growth,  though 
microbes generally have a wider tolerance range than plants. The direction ofN-
transformation  processes  is  dictated  by  the  C: N  ratio  in  the  soil.  A  carbon 
limitation results in net N-mineralization; a nitrogen limitation (for example after 
the addition of excess carbon in the form of organic material with less than 1.3-
PLANT ASSIMILA nON AND N-CYCLING  3 
NZ Fixation 
Nz + NzO 
MineraLization  \  Nitri fication  loenitrification 
..  +  III  /  I 
Organic matter  N H4 •  NO~3up take 
Immobilization  Nitrate reduction  "PLants 
Leachi ng 
Ground water 
Fig.  I.  Principle forms and pathways for  soil  nitrogen (from  Broadbent 7). 
1.5%  N)  results  in  the  net  immobilization  of nitrogen  until  the  C: N  ratio  is 
lowered to 20-30, when net N-mineralization is  reinitiated 8. 
During periods of excess carbon, the free-living N 2-fixers are also favored, and 
fixation  of  considerable  amounts  of  nitrogen  may  occur.  However,  the 
availability of nitrogen to plants during this period is  restricted by competition 
with  the large population of heterotrophs in the soil. 
Under aerobic conditions in most agricultural soils, nitrification is faster than 
nitrate  reduction,  mineralization  faster  than  immobilization, and  nitrification 
NH4 -limited 7. As a consequence, there is often a continual conversion of organic 
nitrogen to nitrate, with little or no accumulation of ammonium. Excess water in 
the  soil  will  inhibit  part of the  soil  aerobic microbial  population, particularly 
fungi  and  actinomycetes.  Also,  aerobic  metabolism  is  more  efficient  for  cell 
synthesis  than  is  anaerobic  metabolism.  Thus,  as  saturation  is  approached, 
immobilization  decreases  faster  than  mineralization,  and  mineral-N 
accumulates 7.  In rice fields fertilized with high rates of ammonium, nitrification 
and denitrification proceed simultaneously: nitrification in the oxidized layer at 
the  soil  surface  and  above,  and  denitrification  below,  where  anaerobiosis  is 
dominant 48.  This is reflected in the better response of rice to deeper application 
of ammonium or urea than to broadcast application 10. 
Nitrogen mineralization proceeds slowly in soils too dry for crops to grow 61 . 
There is also an increase in N-mineralization in re-wetted soil relative to soil kept 
moist 4 ,  and  this may account for  the  initial  flush  of available  nitrogen  in  pot 
experiments or when rain or irrigation follows dry weather when plant growth 
has been restricted. The effects of soil water on microbial activity depend to some 
extent on temperature.  In  one set of incubation studies 13, for  example, rates of 
mineralization dropped by a factor of 3 as soil moisture potential dropped from 
the optimum 0.5 bar (25% H 20) to 2 bar (18% H 20), but the effect of water was 
most marked at the highest and most favorable  temperature, 30°C. In  general, 
low temperature slows down microbial activity, and maximum activity usually 
occurs at ca.  40°C. The period of N-immobilization is  subsequently shorter at 
higher temperatures, although the total amount of nitrogen immobilized is not 
greatly affected 7. 
The surface horizon normally contains nearly all of the mineralizable-N in a 
soil  profile.  However,  there  are  exceptions,  for  example deep  undifferentiated 
alluvial  soils  in which more  than half of the nitrogen  released may come from 
depths below 20cm 13.