• 
• 
Improving Efficiency 
in Calcium Acquisi.t ion 
and Utilization by 
~"""""\ Forage Grasses 
and Legumes 
-
(OlECClON HISTORICA 
_1 
A Proposal ror: Der Bundesminister für Wirtschaftliche 
Zusammenarbeit (BMZ) 
Executing Agency: 
fm Troplca! Agricultura 
Centlo Inlemoclonol de Agricultura Tropical 
• 
s laborating 
tne r: - University of Hohenheim, Stu ttgart , Germany 
542 
.e6 
, I4 J a nllary 1995 
¡ 
IMPROVING EFFICIENC Y IN CALCIUM 
ACQUISITION AND UTILIZATION BY FORAGE 
GRASSES AND L EGUMES 
Special Proj ect Funding 
A Proposal for : BMZ 
1 3 DIC . zoas 
Executing Agency: 
lar Tropical AgrICultura 
de Agricultura Tropical 
Collaborating 
Partner : - lnstitute of Plant Nutrition, University of Hohenheim, 
Stugart, Gennany 
January 1995 
• 
• 
Centro Internacional de Agricultura Tropical CI"'. 
PDO-L003 
January 5, 1995 
Dr. J. de Haas 
Head, Agriculture and Rural Development Division 
Federal Ministry for Economic Cooperation 
and Development (BMZ) 
Friedrich-Ebert-Allee 114-116 
0-53113 Bonn 
Federal RepublicofGermany 
Re!: Propo.mls (or $pedal Projects 
Dear Dr. Haas: 
We are pleased to identify foc youe consideration three special projects involving cooperation 
between CIAT, German partners and NARS: 
(1) Sustainable Animal Production for the. Agropastoral Syst'!ltIS o/ the Brazilian 
Ce"ados 
This propasal was suhmitted in August 1994 and the peer review committee considered it 
to be in principie worthwhile foc funding. We have revised the proposal and have 
addressed tbe tbree issues raised in your previous letter. As per your instructions, we 
have forwarded copies afthe revised propasal directly to ATSAF and GTZ. 
(2) Dynamics and Sustainability 01 Farming and Regional Systems in the South 
American Savannas 
This proposal was submitted in August 1994, but arrived too late to be considered duriog 
the last Special Project Competition. We would appreciate if it would be oow considered 
fo'r the curreat competition. We understand from tbe BMZ fax ofDecember 21 , 1994 tbat 
there is no need to submit additionaI copies oftbe proposal at this stage since there have 
been no changes to the document submitted last August. 
Fil.: PDOJ.19.1 
Mailing address. Apdo. aéreo 671 3. CAl l. COLOMBIA. Telex 05769 CIAT CO. ITT Ol3lcom 10 57 :CG1301 FAX 57-23-647243 . Tel. 57-23-675050 
,. • 
PDO·LOOJ 
(3) Improving Efficiency in Calcium Acquisition amI Utilization by Forage Grasses and 
Legumes 
Copies ofthis proposal have been forwarded directly to ATSAF and GTZ. 
We look forward to continued support for our research cooperation with Gefman 
universities and NARS and we would like to thank you for consideration of eur new requests. 
Yaues sincerely, 
Interim Director General 
Atlachments 
ce: Dr. E. Kurschner, A1SAF 
Dr. J. Friedrichsen, Head ofDivision, Plan! Production, Planl Profeclion and Agricultural ResetÍrch. G1Z 
r PROPOSALS FOR SPECLAL PROJ ECTS El 
Table of Contents 
Lisr oi Figures ......... ..... ..... ..... ................. .. ........ .... ....... . ,., .. ...... .•... ... ...... iv 
Lisl o[ Table;· .. . .. . .................... . .. .. ... .. ....................................... .. . ........... i \' 
Lisl o/ Appendices ..... , .......... . ...... . .......... .... . " .......... , ......... .... ......... ... ", ... iv 
Acrony"'s .... . .. o ••••• " ..... , 00> •• ' .......... . .. . _ .. ............. , .... .. , •••• •• • 0 0 ..... .. .............. v 
1.0 Summary .................. ....... . ..... ... ...... .... ....................... . ....... .... 1 
2.0 Background and Justification ................................................ 4 
2.1 Introduction .......... .. .... . ............. " ................................ " ........... 4 
2.2 Tile target species .... .. ...... ... ....................................................... 5 
2.3 Tlle problem ... ........ .. ... .. ......... .... ............ .... ........ .. ... ........... ...... 5 
2.4 Project outline .......................................................... .............. 1 1 
2.5 Ability of organization and individual s to undertake the research .. . 12 
2.6 Agricultural development and ecology implications of the project. .... 12 
3.0 Project Objectives ..................................... ... .... .. .................. 13 
Goal ....................... ....... " .......... . ............... 0 . . .. .............. . . . . .. ... ...... 13 
Project Purpose .... .. o • • •••• • • ••••• 00_ ••• • ••••••••••••••• • •••••••••• • ••••• • •••• , • ••••• ,., ••• 1 3 
Project outputs .. ..... . .... .. . . ............ . ....... . .................... . .. ... ", . . .... ..... 13 
Benefits for Developing Countries and NARS . ... ... ..... . .. " .. . ....... ......... 13 
4.0 Work Plan . ... ... ... .... ... . ......... .. ........ .................................. ..... 14 
5.0 Training and Workshops ........... ..................... .. . ........... ..... ... 18 
6.0 Expected Patentable Researcb Results ...... .. ...... ... .. . ... .... . ... . 19 
DI 
Table of Contents-Con 'td 
7.0 Funding Requirement s ........................................................ 20 
7.1 Budget .............................. .... ... .............................................. 20 
7.2 Budget notes .. ............................................... ......................... 20 
8.0 References ...................... .... ............ .. ................................... 23 
ID 
List of Figures 
Figure 1: Components of CalciUffi efficiency in plants .... , ........................ 8 
Figure 2: Work Breakdown Structure Linking Project Activities ............ 15 
Figure 3: Project Organization Chart ................................... . .... .. ....... 16 
Figure 4: lmplementation Schedule of Activities ............•.. .. ... . ........ .. ... 17 
List of Tables 
Table 1: Differences in shoot calcium uptake and calcium use efficiency ....... 10 
Table 2: Budget CIAT + Hohenheim University ................... .. ... .. . . .. .... . .... .. 22 
List of Appendices 
Appendix A-1: C.V. Iduputapati M. Rao ........ ....... , ...•.. ,., ... ... ................... 26 
Appelldix A-2: C.V. Horst 1. Marschner .................................................. 30 
Appendix B: Confirmation Partner Letter ........ . ......................... ... ... " .. 33 
Acronyrns 
BM.Z Der Bundesminis ter für Wirtschaftliche Zusammenarbeit, Germany 
(Federal Ministry of Technical Cooperation) 
C IAT Centro Internacional de Agricultura Tropical , Ca li , Colombia 
(International Center for Tropica l Agriculture) 
GTZ Gesse lschaft für Technische Zusammenarbeit , Germany 
(German Agency for Technical Cooperation) 
NARS Naliona l Agricu ltu ral Research Sys lems 
RIEPT Red Internacional de Evaluación de Pastos Tropica les 
( International e lwork for Evaluation of Tropical Pastures) 
.. 
1.0 Surnrnary 
Title: ¡mproving Efficiency in Ca/ci"m ACI/uisition un,1 Utilization by 
Forage Grasses and Legllmes 
Short Title: Calcil/m NI/trition o/ Tropical Forages 
Objeetive of Resea reh: 
Tú identify plant att fibu tes that contr ibute to improved efficiency in 
acquisition and utilizat ion of calcium in Brachiaria and Arac1ús species in 
crder to develop screening procedures to evaluate forage germplasm for 
efliciency in calc iu m acq ui sition and utilization. 
Abstraet: 
The goal oft he Tropical Forages Program al CIAT is to develop forage 
components fo r farm ing systems 00 aeid inferti le soi ls of the humid and 
sub-humid tropics which will contribute to increased and more efficient 
meat and milk product ion , soi l improvement and efosion and weed cont rol. 
Inadequate nutrition is th e main ccnst raint in ruminant productivity 
throughollt most tropical regioos. Developmeot of improved fo rage system s 
of hi gh olltrit ive va lue is the key to iocreased aod sustai oable livestock 
productivity. Forage species can improve soi l fertility , physical structu re, 
aod biological activ ity, protect soi l against erosion, reduce the oeed for 
herbicide lI sed in weed control and sequester large amou nt s of carboo deep 
io the soi l, thus contributing 10 minimizing the green hou se effect of 
iocreasing atmospheric COz' 
D 
lo low fertility acid soils of the tropies, root growth (and partieularly 
rool peoelralion ioto subsoils) is Iimited by the availability of ealcium. 
Forage speeies differ in both requirement io ealcium supply to the rools and 
ealeium demaod per unit tissue weighl. Inter-and intraspecifie differences 
in ealeium efficieoey may be related to a higher effieieoey in utilization 
within the plaot, or a higher effieieoey in aequisitioo of calcium by the 
rools . Furthermore, ¡otracellular compartmeotatioo and concentration of 
calcium io forage tissue affecIs feed value and animal intake . 
The proposed research lo be carried out in Co lombia and Germany will 
test two hypotheses : (i) inter- and intra specific differences in caleium 
aequisitioo are related lo Iheir root arehitecture, cation exchange capacity 
and biodiog of calcium to root cell walls; and (ii) lower co ncentrations of 
calcium in the shoot dry malter resu lt s in lower forage quality . Results 
from Ihis research will help to identify plant attributes that contribute to 
greater acquisition and utilization of calcium in forages . Ident ificati on of 
plant attributes will cootribute to the development of prototype screening 
procedures for tropical forage evaluatioo/ improvement. 
Cooperating Pa rtners: 
(1) Tropical Forages Programo Centro Internacional de Agricultura Tropical 
(CIAT), Cali, Colombia 
(2) Institute of Plant Nutrition, University of Hohenheim, Stuugart, Germaoy 
Names of Principal Scientists: 
CIAr Universily 01 Hohenheim 
• Dr. 1. M. Rae, Praject Leader • Praf. H. Marschner, Project Co-Ieader 
• Dr. C. E. Lascana 
• Dr. 1. W. Miles 
• Dr. P. C. Kerridge 
11 
Staff to be Financed: 
University ofHohenheim CIAr 
• 1 doctoral student (3 years) • 1 technician (2 years) 
• 1 field worker (3 years) 
Budget Total and Breakdown by Activities and Year (Curren! US$): 
The lo/al budget is USS250,520. broken dOWI1 asl ollo\l's: 
CIAT 
Personnel 19,000 20, 100 8,000 47, 100 
Research and Operat ions 18,000 18,000 4,000 40,000 
Training and Workshops 1,000 1,000 3,000 5,000 
lndirect CQsts (20%) 7,600 7,820 3,000 18,420 
Total ClAT 45.600 46.920 18,000 110,520 
Hohenheim University 
Personnel 30,000 30,000 30,000 90,000 
Research and Operations 2,000 10,000 14,000 26,000 
Travel 10,500 8,000 5,500 24,000 
Toral Hohenheim Univ . ..... ___ J__ . .. 42.500 48.000 49,500 140,000 
GRAND TOTAL 88,100 94,920 67,500 250,520 
Au PARTNERS 
D 
2.0 Background and .Justification 
2.1 Introduction 
The avai lab ility and, particularly, the quality of feed 
is the majo constrain t lo increased and more efficient 
animal production in the tropics of Latin America, As ia 
and AfTica. Forage g rasses and ¡egumes can playa role in 
I mtl~e PlllltI\ I tI, I '/ alleviating thi s feed shortage in the majority of 
11m ,mup '(1 In production sys tems (Kerridge, 1994). Further il has been 
( ml,'/J 11m demons trated tha! they have a majar role in contributing 
\11\ I 11 , lo more sus tainable agricultural sys tems (Vera el al. , 
1992). The combinalion of nitragen fixing rorage 
" I 
¡egumes wit h deep- rooted grass can ¡ncrease nutrient 
cycJing, g reatly improve animal production, markedly 
in crease soil b io logical aClivity , and s lo re significant 
amounts of o rganie earbon deep in the so il (Thomas et 
al. , 1994; Rao el al., 1994; Fisher el al. , 1994). 
Low nutrient supply is a major limitation of forage 
adaptat ion and produclion in acid soils of Ihe tropies . The 
lI se of forages adapted to low fertility acid soil s is one of 
the most effeetive means o f managing these soils . 
Considerab le aehievements have been made in identifying 
legumes and g rasses we ll adapted lo these so il s buI little 
is known abou t the meehani sms for adaptat ion (Rao et al. . 
1993). Widespread adoption o f forage cultivars depends 
on e ffi eient aequi s ition of nutrients from the soi l and 
utili zation for g rowth . 
"ir/. 'pf(!tUl ttd()l'flIIII " 
/ora;:(' ( ,,¡tú'.,n If('pi ,ul, , Continued progress in the se leetion and genelie 
tI" ir fI" ¡en, lH //1'; ,il/vll o{ improvement o f forages will depend upon the 
tl/ttri e 11 h ./mm 11/11 lI.:J" 111 la ",h development of rapid and reliab le teehniques whieh 
facilitate sereening of large numbers of genolypes fo r 
toleranee lo low fertility aeid soil s . 
a 
H\(h. ROL'!) \\nJl~IPi( \110\ 
2.2 The target species 
The genus Brachiaria is the source oC a number of 
widely used tropical forage grasses. Together, several 
JJrlllfti"ria 'I'C{'I('\ lit( IJ¡ Brachiaria species are undoubtedly the most widely sown 
mmt II'jdl'~r \011'11 IjlnI~ \ If forages in tropical America, with 40-50 million hect ares 
tropical llt/aÍl II sown in Brazil alone. An important germplasm collection 
was assembled al CIAT and regional evaluation of Ihis 
coll ection is essentially complete. A genetic 
enhancement project is in progress at CIAT which utilizes 
new techniques in exploiting apomixis to create synthetic 
gene pools with res is tance to major pests (spittleburg and 
leaf cutter ants), high feed quality and good persistence 
on acid, low fertil ity soi ls. 
Arachis pinto ; is unique among tropical rorage 
legumes in the humid tropics in its ability to form stable 
grass- Iegume associations with vigorous grasses like the 
Brachiarias. It is persistent and even ¡ncreases under 
heavy and continuous grazing pressure. Nutritive quality 
Ira,",,;, [linIo; l\ II,.'II-udl/prl ti 
is high. However, the present widely used cultivar CIAT 
lo I//wjali/ily add wi/, 
17434 is slow in establishment and is not tolerant of dry 
conditions. Research efforts are directed towards 
increas ing the range of adaptation. 
2.3 Tbe problem 
In many tropical soils, low calcium (Ca) may be a 
grea ter limitation to plant growth than high aluminum 
(Al) buI has received much less attention. In highly 
weathered tropical soils (Oxisols and Ultisols), plant 
growth is often Iimited by Al toxicity and phosphorus and 
Ca defi ciency (Marschner, 199 1). In the subsoil . the Ca 
content i5 often ve ry low. Many subsoils have less than 4 
mg Ca kg·1 ofsoil (Ritchey el aL , 1987). Because Ca is 
/" '1/;:"11 H'l (Jlllt,.,t/ trtJ/" I nol mobile in the phloem it do es nOI move downwards in 
\oil\, ("h'lIIm "ut Uf 111 1ft Ihe root5 towards the root tips where it is required for 
,,,h,¡Ji/, ;\ ,~frf!1I \'(.'1)' (011 growth (Hanson, 1984). Thus, apical root zones (rool 
tips) have to cover their Ca demand for growth by direct 
uptake from their immediate environment. In acid mineral 
soils, root growth is Iherefore often severely limited, 
particularly root penetration into the subsoil (Pearson et 
D 
BACKGROUND ANO JUSTlFlCATlON 
al. . 1973) and , correspondingly, utilization of nutrients 
and water from ¡he subsoi l (Howard and Adams, 1965). 
Lo\V Ca activity in the s ubsoi l may be a greate r limitation 
lo plan! growth than high Al (Bruce e l aL , 1988) . 
(JI AnOlher very importan! aspecl of Ca nutritíon is the 
AIt importan! aspect effect of Ca concentratíon on feed va lue and animal 
(,(rlá"", nlltrilion o.{trop;clIl intake. Calcium fed as a supplement is nol as effective as 
f(}rages is the elfecl o/ higher Ca in the plan! tissue in improving fo rage quality 
calciulU CONcentralion 011 (Minson, 1990). Therefore \Ve need to identify forage 
forttge qflalit)' species and ecotypes that can gro\\' and effici ently acquire 
Ca from soil s very low in the nutri ent. 
Calciul11 nutrition is different 10 other nutrienls 
(Marschner, 1974). Because of its phloem·immobility 
there is no re·distribution of Ca. 11) root 10 shoot t ransport 
and distribution within Ihe ShOOI, Ca is confined lo the 
xylem and , thus , is closely related lO Ihe Iranspiration 
slream . The Ca transport into low transpiring tissues such 
as shoot apices and young leaves is therefore low. In 
addition , Ca binding to pectales in cel! walls or 
precipitatíon as oxalate along the transport palhway 
furt her limil Ca supply lo Ihe s ites of demand for growth . 
Furlhe rmore, the common route for recycling nutrients in 
the ph loem from mature leaves to new growth is nOI 
availab le for Ca (Ho and Adams, 1989). In vigorous 
growing trop ica l forage grasses and legumes, un less Ihe 
import of Ca vía xylem matches the demand during Ihe 
acceleraled expansion growth for new leaves, plant 
growlh will be limited by Ca su pply. Recent fie ld 
expe riments with tropical fo rage grasses (Brachiaria 
species) on Oxisols in Carimagua (Colombia) s upport fhi s 
view of Ca limitation. Increase in shoot dry matter 
production was similar wilh Ca app lication in form of 
lime or gypsum, although so il pH was only increased 
with lime (K. Haussler, unpublished res ult s). 
There is variar ion belween planls in their abil ity 10 
g row on low Ca soi1s. Plant species differ in both 
Plmrf species d~{fer in fhe;r requirement in Ca supply to Ihe roots and Ca demand per 
ahility 10 acqu;re (lfl(luti/ize unit ti ssue weight. As a rule, Ihe requi red slipply (i.e .. 
calciumfrom soits so il content or concentrat ion in Ihe so1 1 solution) of Ca 
and the Ca content in the ti ss ll e are higher in 
11 
n \( h.(,f(Ol '\D .\I\D·II ~11F f( \ 110\ 
dicotyledonous species (e.g., legumes) Ihan in 
monoeotyledonous species such as forage g rasses 
(Loneragan et al. , 1968 ; Loneragan and Snowball, 1969; 
Is lam et al. , 1987). But also between species of grasses 
and dicots as well as between cultivars or ecotypes of a 
given species considerable differences exist in the 
apparent requirement and demand of Ca. According 10 
Ihese differences the lerms " calcium-efficiency" and 
"calcium-ineffic iency" have been introduced. From the 
agronomical or breeding poinl of view, for adaptation lo 
acid mineral soil s a high calcium-efficiency is achieved, 
i.e., genotypes which acquire grea ter amounts of Ca and 
produce a high yie ld in a soi l low in avai lable Ca. 
J)~l.Ierf.·ltn'\ in caldum Calcium effíciency may be related 10 a higher 
efficicl1(r muy he rt'latct! lo effíciency in ulili zat ion within the plant , or a higher 
effi ciency in acquisilion of Ca by the roots (Figure 1). 
ucqui\ifilll1 hy roO/'i, OT 
lar Examples for a more effí cient utilization of Ca wi thin the 
11 ,ili:.aúo/l p lUI11 groll'(h 
sheot have be en given for a Ca-effícient tomato cultivar 
(English and Barker, 1987) or cowpea cultivar (Horst el 
al. , 1992), o r by a hi gher proportion of Ca translocated lo 
the shoot apex in a Ca-effícient tomato cultivar (Behling 
et al., 1989). Differences in efficiency in Ca acquisition 
between species and genotypes within a species are well 
documented and demonstrated by differences in Ca 
concentrations in the shoots of plants grown in the same 
solution (Loneragan et aL, 1968; Islam el al., 1987) or the 
same soil. 
Inter- and intraspeci fí c variation in Ca effici ency has 
been demonstrated in several acid soil-adapted tropical 
forage grasses and legumes in fí eld experiments 
conducted on an Oxisol of Carimagua (Colombia) 
(Tropical Pastures Program Annual Report , 198 1). 
1'111' hig!tcr t{fiál'It(T ¡" Among the grasses, Brachiaria hl/midicola CIAT 679 was 
(:tll('/II111 U('(II/hitioll 11'1/\ the most Ca-effícient one, il had the lowest requirement 
relall,tI lo Iti;:hl..'r mol for Ca supply (125 kg CaCO, ha"' ) and also the lowest 
Imlll! hi,,;.: internal Ca concentration (0 .22% Ca in the dry matter) . 
Glasshouse studies using acid soi ls of conlrasting tex ture 
have indicated marked interspecific va riation in Ca 
acquisition and internal utilization among 7 grasses and 
12 forage ¡egumes (Table 1; I.M . Rao et al. , unpubli shed 
data) . Recent work, supported in a BMZ project, 
a 
Figure 1 
Components of calcium efficiency in plants 
Calcium Efficiency 
Efficiency in Utilization 
• Low demand of calcium at • High root-to-shoot transport of 
cellular level calcium 
• Preterentlal transport of calcium 
• Maintenance of high concentra~ Efficiency in Acquisition to the shoot apex and young 
tions of soluble calcjum leaves 
Root physi%gy 
Rool morpho/ogy • High affinity uptake system for 
calcium 
• Large surface area 
• Root exudates 
• Abundant root tips (complexation of aluminum) 
-8-
demonstrated higher Ca acquisition in Brachiaria 
rllziz;ens;s compared to Brachiaria dictyonellra, leading 
to much higher Ca concentrations in the shoot dry matter 
and uptake of Ca ln the above·ground biomass of B. 
ruziziensis (K. Hauss ler, GTZ Report , 1994). The higher 
efficiency in Ca acquisition in B. ruziziensis was re lated 
not to a higher root length but higher branching, i.e., more 
root tips which are the main si tes of Ca uptake along the 
root axis (Marschner and Richter, 1974; Haussling et aL, 
1988). 
Much of the 50 million hect ares planted to Brachiaria 
species pastures in tropical America are now in a state of 
degradation owing to nutrien t depletion. There is good 
evidence that inclusion of legumes can assist in reversing 
pasture degradation. Arachis pinto; forms stable grass· 
rhC'r( i, l/O hIU1H'lctlJ:c 1111 legume associations with the Brachiaria species. But 
('('o(¡pit" diJlerem:c,\ ;/1 there is no information on differences in Ca acquisition 
("/lIC;1I111 (/("{jlú,iriofl (1/1(/ and utilization among different A. pintoi ecotypes now 
IItilbllhm hy .1radm piut(l; available . A. pintoi is better adapted to c1ay loam than 
sandy loam Oxisols which may be related to better water 
and Ca availability in the clay loam. 
The Ca concentration per unit shoot dry weight is 
much higher in A. pinto; (dicot) compared to the forage 
grasses. In preliminary experiments with A. pintoi grolJJ!l 
in two Oxi sol s (sandy loam, c1ay loam) the Ca 
concentration in the shoot dry matter was about 15 mg g.l 
dry matter (Milz, unpublished) . So far only a few data are 
available on Ca acquisition and internal utilization in a 
single ecotype, CIAT 17434 , of A. pintoi (See Table 1; 
LM. Rao et al. , unpubli shed res ults). Shoot Ca uptake , at 
high Ca supply. was greater with c1ay loam than sandy 
loam soi l. 
Agronomic evaluation of thirty·three ecotypes of A. 
';.:rollomic c l'tduurú", iu lile 
p;nto; in the Cerrados of Braz il indicated ecotypic 
Cl'rrmlm al' /Jm:.iI ¡m/j{"(//(·tI 
differences in tolerance to so il water deficits . Based on 
(!(''''-I,/Iic I'tlriatitm in ti,...\' green leaf retention in the dry season. the performance of 
\etnOIl /lt'l:for!wlIlce oI CIAT 22160 was markedly superior lO CIAT 17434 (E . 
.· ¡rach;" f1intm iza rro, unpubli shed result s). But there is no information 
on differences in Ca acquisition and utilization among A . 
pintoi ecotypes. 
Table 1 
Differences in shoot calcium uptake and calcium use efficiency in Brachiaria and Arachis 
species grown in low fertility sandy loam acid soil. 
Species and Ecotype Shoot Ca uptake (mg/pot) Ca use efficiency (g/g)* 
Brachiaria decumbens 20.1 211 
CIAT606 
Brachiaria brizantha 19.9 170 
CIAT6780 
Brachiaria dictyoneura 11 .5 187 
CIAT6333 
Brachiaria humidicola 9.5 187 
CIAT6369 
Arachis pintoi 60.5 34 
CIAT 17434 
• grams of forage produced per gram of total Ca uptake from soil 
_r. .. _. ... 
, .10 ~00\lC' __ 
·10-
11-\( Jo.:(JItOI '\1) \ U 1( I H I( \ 110 
2.4 Project outline 
We intend to test two hYPolheses : 
(n Tha! differences in Ca acqusition among and within 
Brachiaria and Arachis speeies and eeo~'pes are related to 
thei r root arehiteeture, cation exchange capacily and 
binding of Ca to root cell walls, 
(1) Thal lo",er Ca conccntrations in the shool dry matter affeet 
forage quality, 
By testing th ese hypotheses and identifying plant 
1m/Ir /q ti \( r('t fII attribules that contribute to greater acquisition and 
"me, dllrn will "o.; ti 1/ utilizalion of Ca, we ShOll ld be able lo develop improved 
Jár Ir¡ll'intl/(lfIl~(, Id/llOft sc reening procedures for tropical forage evaluatíonl 
amI iml'rll1'CI1I('"1 improvement. Higher Ca acq ui si tion will enhance 
recycling of Ca in tropical pastures and reduce Ca losses 
from Ihe system by leaching, 
Grasses and legumes are known to contri bu te not 
only 10 animal prodllction buI al so 10 soil ferril ity in 
tropical farming sys tems, ThllS they are essential 
components of sustainability. 
What is desi rable from an animal nutrition poin! of 
view is Ihe greater efficency in acqu isition of Ca by rools 
and translocation 10 the shoots. There is a need to relate 
f om;:l' grt/\\t'\ (1m! /t'glllll \ the effi c ieney of Ca acqllis iti on to Ca aetivity in so il 
(Ir" ('\\('III;t/1 "Olll/Ulm 11/\ III solution. This project will con lribllte to se lection of high 
ugrh U/fura! \/I\lt,;/wbilifr qllality forages and will a1so have an addilional spin-off 
on fertilizer management for pastures on low fenility aeid 
soil s. By re lating Ca acq uis ition to so il Ca jt wil1 provide 
important information to g rowth of other erops on aeid 
soi ls , 
m 
2.5 Ability of organization and individuals to 
undertake the resea rch . 
(,'cr/1/all (Hui 'm al IllliJ'('r\¡fic\ The proj ect's principal CIAT scientis ts Drs . I. M, 
11';11 ('o"uhortllt' ¡" ,Il(' Im~i('{'( Rao (Project leader) , C. E. Lascano, J. W, Miles, and P. 
C. Kerridge have long- term experience in the 
deve lopment of tropical rorage germ plasm, namely in the 
areas of plant nutrition/physiology, forage quality/animal 
nutrition, forage geneti cslbreeding, and soil fertil ityl 
ag ronomy, res pectively. They will be ab le to provide the 
required local supervis ion to the German doctoral student 
and the eventual B. Sc, students. 
For the past 3 years , th e CIAT Tropical Forages 
Prog ram has been cooperat ing c10sely with the German 
proj ec t partner (Prof. H. Marschner) wi th special proj ect 
fund ing from BMZ in a project entitled " Study of 
phosphorus dynamics in the rhizosphere of various grass 
and legume species g rowing in acid soi ls of Lat ín 
Ameri ca", 
2.6 Agricultural development and ecology 
implications of the project 
CukiuIII cjfiÓ('lIlfortlgn ,,'íll Development of calcium effi cient forages Ibat 
nmtrih"fc lo \u\laitrah!(' combine productivity and qual ity with adaptation to low 
/i1'(!.\lod. . produdion ferti lity ac id soi ls of the tropics, will contribute to 
sustainable lives tock production. 
In addi tion, the calcium effi cient [orages will 
enhance recycling of calcium in agropastoral systems, 
reduce calcium losses from the sys tem by leaching, and 
contr ibu te to sustainable land use sys tems in the humid 
and subhumid tropics. 
m 
3.0 Projcct Ohjecthcs 
Goal 
To develop tropica l forages w ith improved efficiency 
in acquisi ti on and utilization o f calcium from low fertility 
¡-"" acid 50i1 5 . 
prO;j'('( It{/\ I ,",ca!' J.: //1 
II'/rit" (omph /11 ,,'\ mld 
IIIh'J;rtlln ¡"t,) thr r \( a I Project Purpose 
,Ji( (1 j r I rtJ/UI.:al lor 
Pro.", I To ident ify plant attributes tha! contribule 10 
improved efficiency in acquisi tion and utili zation of 
calcium in Brachiaria and Arachis in order to develop 
sc reening procedures to eva luate forage ge rmplasm for 
efficiency in caJcium acquisition and utilization. 
Project outputs 
The expected outputs from the aboye objectives are: 
.. ldentification of 8rachiaria and Arachis species and 
ecotypes with greater ability to acquire and utilize calcium. 
I 111' (~'l'('¡;(('tI OIlIPIl/\ 1I If • Identification of plant attributes that cont ribute to calcium 
he/" lo Idellfify caloll", efficiency in Brachiaria and Arachis. 
(~flh ir", frol'l('u!Jord,. • Selection of superior Brachiaria genepools (using new 
grm ., (/111111);1111"\ sc reening metholodology). 
• Benefits rOr DeveloJling Countries and NARS 
The results from the project wi ll be presented at 
reg ional conferences organized by regional networks such 
as RlEPT and Agropastora l Ne twork . Furthermore, fina l 
result s of the project wi ll be di scussed in a 2·day 
workshop at CIA T. 
The outputs of thi s project wi ll contribute towards 
the deve lopment of forage components for farming 
systems on acid infertile soil s of the humid and sub­
humid tropics which is the broader goa l of the Tropical 
Forages Program at CIAT. 
m 
4.0 Work Plan 
The principal activities and subacti vi ti es of the 
project as they relate to the project's outputs are 
illus trated in Fig ure 2 . The project organization 
concerning technical reporting and financia! management 
is shown in Figure 3 . Figure 4 shows the implementation 
schedule with the cornmencement and duration of each 
main activity. 
The execution of the project in PalmiTa and 
eaTimag ua, Colombia will require th ree years . The 
project will be coordinated by Dr. L M . Rao, Tropical 
The prtljl'( t lI'ill h Forages P rogram, e IAT, with assistance from o th er 
(,f1ordillflfl''¡ by ,he I rfll';j 11/ scienti s ts from CIAT and Pref. H. Marschner from the 
I úraJ.:l',\ ProJ.:ra",. (lItd 1»0,11 University of Hohenhe im. Germany. 
(H'rrlltlll ami (olm"hilftl 
The research will be carried o ut by a Ph. D. s tudent 
\fllt/('IIf\ flm!.\óenr¡\!\ 1 ill 
and staff of Ge rman University and CIAT scienti sts (see 
('I/rr.r oul ,he 1'C\'can" 
al so 5.0 Training) 
Figure 2 shows the breakdown of the project 
s tructure with desc riptions o f the goal, purpose. o utputs 
and activities . 
m 
Figure 2 
Project Description 
Struclure Unking Projecl Activlties lo Projecl OUlpUls 
Program Goal 
T o develop tropical lorages with improved efficiency in 
acquisition and utilization of calcium from low fertility acid 
soils. 
Project Purpose 
T o idenlify planl atlributes that conlribule to improved 
efficiency in acquisition and utilization of calcium in 
Brachiaria and Arachis in order to develop screening 
procedures to evaluale lorage gennplasm lor efficiency in 
calcium acquisition and utilization. 
Identification of Brachiaria and laentification of plant attributes Selection of superior Project management 
Arachis species and ecotypes that contribute to calcium Brachiaria genepools 
with greater ability to acquire efficiency in 8rachiaria and (using new screening and dissemination of 
Arachis results 
Select contrasting geootypes of SekM:t conlra5ting genotypes al SfleN.". In<! Grow out gene pools (advanced Revise proposal together with 
Brachiaria and Arachis trom agronomic Ar~ch/s ffom agronomle evalultion. fines) from Brachlaria donor. 
evaluation experiments. Conduet I glasshouM experIment with $aOCIy loan acicl improvement programo 
subsoll supplied with difl"erent ...... and fom\S (lime, 
Establish an experiment on a calcium gypsum) al Ca, delenniM rOOl atehitecture using Provide progress, technical and 
deficieot sandy Ioam acid subsol with mlnimizotrons In<! measure Ca COOC9Otratlom in rOOl Undertake restricted sel of financial reports. 
different levels of Ca supp/y as ~me and and shoots. measurements on plant attribules 
gypsum. UH of labeIIed isotope (""'Ca) lechnique 100 as would be usad by a Geoeticist Report Pfogress at Regional 
measurement of Ca uptake in difl"orent root ¡:OO«! screening material. Networ1< meetings. 
Determine dry matter partitioning among (mature, eIongaling, lip) of pómary and secondary 
plant parts and leaf area production (three rooI$. and of C. lransport lo lhe shooI:s and young 
leveIs in particular. Validate laboralory findings 'Nith Publish results. 
times during grov.;ng season), 
Deteonlne shotHerm uptake of calcium ("'Ca) in roots restricted set of animal feeding 
concentration of Ca and another mineral exposed lo soIutions of differenl pH Ind eoneentratlons trials. 
elements in the dry matter. Conduct a workshop al eIAT. 
of I luminum (water Ind $IInd cultur. .xperiments). 
Determine root biomass and root length DeteoniM cation exchange capaeity fA lhe c.M waUs in 
roots and e.tI waU constituerts .nd digestibility of lear Select Ca efficieot Brachiaria Provide final reporto 
distribution across the soil profile (up to and stom tlssue. gene pools. 
80 cm soil depth). Determine Intraee!lular eompartmentatlon and binding 
Determine partitioning of nitrogen among fom1S fA ca In laavM of dílferent age. 
Determine eell wall comtituent:s and digastibility of leal" "' ..... ',. .. _'..  -
soluble and total protein in leaves of ,. . ~""""" ..- -
and stem tissues. 
different maturities. Relala compartmentatlon In<! blndlng forms d ca with 
Identify Ca-efficient 8rachiaria and Iorage quallly attributes. 
Arachís ecotypes. o.v.Iop • ~ seraening proce<tura 10 evaluale Ca­ -15-
efflcleney. 
Relalo C. acqubitJon to Ca activily in soil soIllIlon. 
Figure 3 
Project Organization Chart 
~----
BMZ GTZ ] 
CIAT Offie. 01 
Director General 
,----­ CIAT Offie. 01 Deputy CIAT Deputy Director of 
Director General/Research Finance and Administration 
I 
I 
I 
I 
I ,J,---,. 
CIAT Research - ------' Tropical Forages CIAT Projecl 
Project Coordinator Program Leader Support orfice 
L _________ _ ____________ __ ___ _ _ 
~~~~- I~~L-~ 
National University, University of Financia! Analysis Monitoringl 
Palmira, Colombia Hohenheim and Reporting Evaluation 
Research 81aft ) 
Communication lines - - - - - - -
Reporting Unes 
--­ -16-
,,~ 
Figura 4 
Project Implementation Schedule of Activities by Quarter 
Yea!' I Year 2 Year 3 
Acth<ities 
Q1 I Q2 I Q3 I Q4 Q1 I Q2 I Q3 I Q4 Q1 I Q2 I Q3 I Q4 
IOENTIFICATION OF E COTYPES 
Selection 01 ecotypes .. .. .............. .. .. .. .... ..... ..... ........ .. .. . ............... .. ..  
I 
Establishment 01 field experiment . .... .. ....... .. .... 
Evaluation 01 shool and root produclion ... ... ..... ..... ....... ......... ... ...... . .... . .. 
I 
Determination 01 Ca efficiency ............ .... .... . ... ..... .. .. 
I 
Identifica non 01 Ca efficient ecotypes ......... ........ ......... ...... .. ...... .. .... .. . .... . . .. 
IDENTIFICATION OF PLANT A TTRIBUTES 
Selecl contrasting ecotypes .. ...... ..... . ... ... ... ...... .................. .. .. ..... . ... 
I I 
Eslablish glasshouse experiments .. .... .... ... ...... ....... ... .... ..  
I 
Isolope sludies on Ca acquisition .... ........ ........ .. .. .. ..  
I 
Studies on forage quality and Ca compartmentation .... ..... .. .. .. ........ 
I 
Screening procedure lor Ca efficiency .. .. ........ .. 
SELECTION OF BRACHIARIA GENE P OOLS 
EstabHsh field trial ....... ... ......... .... . .................. ... .. .. .. .. ....... ....... .. .. ..... .. .. ..... .. ............ 
Test screening procedures . ... ....... ..... ..... .... ... .. .. .... . .. .... .. . .... ........ .. . . ..... .... ......... .. 
P ROJECT MANAGEMENT 
Workshop ..... .. ... .... ..... ... .. .... .. .. .... .... ... .. .. .... .. ........ ..... ..... ... .. ... ... ... .. .. ..  
Publication 01 results .................. .. ............................ .. ..  ~ 
'_I>._~ 
......... ""'Ñ'-' - \ 7-
5.0 Training and Workshops 
A German docto ral sl udent frcm the Univers ity of 
Hohenheim wiU carry out Ihe proposed resea rch. He will 
work under Ihe supervision of Ihe project leaders, Dr. I. 
Sllit!enr~ from che local M. Rae (CIAT) and Praf. H. Marsch ner (Universi ly of 
IIniversil)' wíll he gh'en Ihe Hohenheim) . 
opportllllir)' lor 
IIndergrtltlllllle thesi.s rc.w:nrclt Th e projecl will also provide Ihe opport unity for two 
Colombian unde rgrad uate s tud en ts frcm rhe University al 
PalmiTa (Uni versidad Nacional) , Colombia . According lo 
jls gender paliey, CIAT will ta ke ca re tha! bolh female 
and olaJe students wilt participate in Ihe project if Ihey 
are equall y tal ented. 
A Iwo-day workshop wil! be conducted a l CIAT al 
Ihe end of Ihe p roject 10 discuss final resu lls and Ihei r 
implica tions for fulure research on the deve lopment of 
forage components for s ll s tainab le agro pastoral systems 
on low rert ility ac id so ils of the humi d and sllb-humid 
trop ics. 
El 
6.0 Expected Pateníable Research Results 
CIAT endorses the principi e of free access 10 
research result s. 11 supports th is through publica tion of 
research findings in international journals and in-house 
documents . 
There are no patentable res uhs anticipated in Ih is 
project. 
m 
I 
I 
7.0 Funding Requirements 
I 
I 
7.1 Budget 
I 
The budgets for CIAT and the University of 
I Hohenheim appear in Table 2. The CIAT budget will be 
admini stered by e JAT. 
7.2 Budget notes 
Personnel 
The two pos itions (1 techn ic ian and 1 worker) 
budgeted for CIAT refer lO assis lance needed for one 
technician al CIAT-Carimagua for fíe ld research and one 
worker al CIAT · PalmiTa for glasshouse resea rch. 
The German doctoral student will ca rry out Ihe 
proposed research with Ihe help of a technician and a 
worker al CIAT (18 months) and al Ihe Univers ity of 
Hohenheim ( 18 months). The doctora l student wi ll be 
supported by tWQ Colombian undergraduate students . 
Research and operational expenses 
As the project ¡neludes fie ld and glasshouse resea rch, 
there are substan tial cos ts ¡nvolved in the establishment 
of experi menls. purchase of chemicals including 
radioi sotopes. process ing of samples and mineral analys is 
of samples . 
Research results from the project w ill be published in 
three languages (Eng li sh. German and Spani sh) . 
r_,....., ..,. __ .""~ ........... ,.,~, .. _ El 
Training "nd Workshops (CIA T budget) 
• Provision is made in the projccI [or trainins up to two 
B.Sc. snldenlS from a Colombian Uni"crsity whe "ill 
receive a 5mall alJowance to do undergraduate thesis 
research . 
• The workshop budgeted refer to a short (two-day) meeting 
al CJAT \\ith the participat ion of CIAT scient ists, the 
Gcnnan doctoral student. and Ihe sc ientists from NARS al 
Ihe end of lhe project. in arder 10 discuss final results and 
their implications for futmc rescarch. 
Tra"el (Hohcnheim budget) 
The amOlln t budgeted for local ¡ravel refers to an 
absolute minimum necessary for Ihe doctoral st udent lo 
move between CIAT-Palmira and CJAT-Ca rimagua. 
The international travel [or lhe doc to ra l s tudent and 
supervisors ¡neludes a provision [o r an eventual 
participation in an inte rnational congress . The German 
supervisor wi ll travel to Colombia dllring Year 1 10 visil 
field and g lasshollse experimenls and 10 discllss the 
project wi th CIAT sc ienlists. The C IAT s upervisor will 
Iravel to Germnny al lhe end of Year 2 10 discuss Ihe 
progress mad e. 
m 
Table 2 
CENTRO INTERNACIONAL DE AGRICULTURA TROPICAL - CIAT 
BMZ - Improvlng efficlency In calcium acquisilion and utili zatlon by foraga gresses and legumes 
Propasad Budgel 
(IN US$ ) 
INSTITUTION/LlNE ITEM Vear Vear 2 Year 3 Total 
CIAT 
Per90nnel 
Techniclan "} 12,000 12,600 24,600 
Waker (1) 7,000 7,500 8,000 22500 
Tolal personnel 19,000 20,100 8,000 47,100 
Research and Operation s 
Supplies 8'ld S8Mces 12,000 10,000 22000 
Labcratcxy Arl~sis 6,000 8,000 4,000 18,000 
Total ressarch and oper8tions 18,000 18,000 4,000 40,000 
Training and workshops 
B.Sc. studerlts (2) 1,000 1,000 ~OOO 
Workst1ops 3,000 3,000 
Tolal tralning and workshops 1,000 1,000 3,000 5,000 
Indlrect costs (20%) 7,600 7,820 3,000 18,420 
TOTALCIAT 45,600 46,920 18,000 110,520 
Hohenheim University 
Personnel 
Doctoral student (1) 30,000 30,000 30,000 90,000 
Total personnel 30,000 30,000 30,000 90,000 
Research and Operations 
Supplles 2,000 8,000 I ~OOO 22000 
Publcations 2,000 ~OOO 4,000 
Total research and operalions 2,000 10,000 14,000 26,000 
Travel 
Local (PhD. Student) 3,000 3,000 3,000 9,000 
InternaticnaJ ( PhO. suden! ) 2,500 ~5OO 5,000 
(SupilM9Of ) 5,000 5,000 10,000 
Total travel 10,500 8,000 5,500 24,000 
TOTAL HOHENHEIM UNIVERSITV 42.500 48,000 49,500 140.000 
GRANO TOTAL 88,100 94,920 67,500 250,520 
--_~-..  
J?UMfk<¡f'"""-<-ú, ,'  
Juan A. Garafulic 
Financlal Controller 
8.0 References 
Behling, J.P ., Gabelman, W.H. and Gerloff, G.C. (1989) . The distribution and utilization 
of calcium by two tomato (Lycopersicon esc"lenlllm Mili.) Iines differing in 
calcium efficiency when g rown under low·Ca stress. Plant SoO 113 , 189-196. 
Bruce, R.e., Warrell, L.A., Edwards , D.G., and Bell , L.e. (1988). Effects of aluminium 
and calcium in the soi l so lution of acid soils on roOl elongation of Glyc;ne max (L.) 
Merr. Allst, J . Agric. Res . 39, 319-338 . 
English, lE. and Barker, A.V. (1987). Ion interactions in ca lcium-efficient and calcium­
inefficient tomato lines. J. Plant NlIlr. 10, 857-869. 
Fisher, M.J ., Rao , I.M ., Ayarza, M.A. , Lascano , C.E. , Sanz, J. I., Thomas, RJ. and Vera, 
R. (1994). Carbon storage by introduced deep-rooted grasses in the south American 
savannas. Na llo'e 371 : 236-238. 
HauI31ing, M., Jorns, C.A., Lehmbecker, G., Hecht-Buchholz, Ch. and Marschner, H. 
(1988) . Ion and water uptake in relation to rOol development in Norway spruce 
(Picea abies (L.) Karst .) . J. Plan! Physiol. 133,486-491. 
Hanson, J .B. (1984) . The functions of calcium in plant nulrition . Advances in Plant 
Nutrilion . Vol. 1, 149-208. 
Ho, L.C. and Adams, P. (1989). Calcium deficiency-a matter of inadequale transport to 
rapidily growing organs. Plams Today. pp. 202-207. 
Horst, W.J. , Currle, C. and Wissemeier, A.H. (1992). Differences in calcium efficiency 
between cowpea (Vigna IInglliclllara (L.) Walp.) cultivars. Plant Soi! 146, 45-54 . 
Howard, 0 .0 . and Adams, R. (1965). Calcium requirement for penetration ofsubsoil by 
primary cotton roots . Soil Sci. Soco Am. Proc. 29, 558-562 . 
Islam, A.K.M.S., Asher, C.J. and Edwards , D.G. (1 987). Response of plants to calcium 
concentration in flowing solution culture with chloride or s ulphate as the counter­
ion. Plan! Soi/98, 377-395 . 
Kerridge, P .C. (1994) . Opportunities for forage research and development in tropical 
Latin America. In: D.F. Cameron (Ed.) Strategic Directions for Tropical Forage 
Research in ACIAR . ACJAR Proceedings, Canberra, Australia (in press) . 
m 
REFERENCES 
Loneragan, J.F. and Snowball , K. {l969) . . Rate of calcium ab sorption by planl roots and 
il s relation lo growth . AUS1. J . Ag l'ic. Res. 20, 479-490 . 
Loneragan, J.F., Snowball , K. and Simmons, W.J. (1968). Response of plants to caleium 
eoneentration in solution culture. Alfs. J. Agric. Res. 19, 845-857. 
Marschner, H. (1974). Calcium nutrition in higher plan ts. Ne rh. J. Agric. Sci. 22, 275-282 . 
Marschner, H. and Riehter, C. (\ 974) . CaJcium-Transport in Wurzeln von Mai s- und 
Bohnenkeimpilanzen. Plal1l So;! 40, 193-2 10. 
Marschner, H. (1 99 1). Mechanisms of adaptat ion of plants to ae id soi ls. Plam and So ;/ 
134. 1-20. 
Minson, D. J . (1990) Forage in ruminant nulrition. Academic Press, San Diego. 
Pearson, R.W., Childs, J . and Lund, Z.F. (1 973). Uniformity of Iimestone mixing in aeíd 
subsoil as a factor in cotton rool penetralion. So il Sc;' Soco Am. Proc. 37, 727-732 . 
Rao, I.M ., Zeigler, R.S., Vera R. and Sarkarung, S. (1993). Seleetion and breeding for 
acid-soil tolerance in crops : Upland rice and tropical forages as case studies. 
BioScience 43 : 454-465 . 
Rao, LM ., Ayarza, M.A. and Thomas, R.J . (1994) The use ofearbon isotope ralios to 
evaluate legume cont ribution to soi l enhancement in tropi cal pastures. Plam and 
So il/62 : 177-1 82 . 
Ritchey, K.O., Sousa, O.M.G. and Silva, J.E. ( 1987). Calcium and rOOI penet ration in 
highly wathered soils. In: L.M. Gourley and J .G. Salinas (Eds .). Sorgl11ll11 for Acid 
Sol/s. CIAT, Cali, Colombia, pp. 11 7- 132. 
Thomas, R.J ., fisher, M.J. , Ayarza, M.A. and Sanz, J.1. (\994). The role of forage grass 
and legumes in maintaining Ihe productivity of acid soils in Latin America. Adv. 
So il Sei., in press. 
TPP Annual Report . (198 1) . Tropical Pastures Program Annual Report . CIAT 
02ETP( 1) 82. September, 1982. CIAT, Cali , Colombia. pp. 304. 
Vera , R.R., Thomas, R.J., Sanint, L. and Sanz, J .1. (1992). Development of sus tainable 
ley-farming systems for the aeid-soil savannas of tropical America. Anais da 
Academia Brasileira de Cienc;a s 64: 105-125 . 
m 
Appendix A-1 
c.v. 
Idupulapati M. Rao 
Position in Project: Research Coordinator, Plant NutritionistlPhysiologist. 
Intemational Research 
Experience: Centro Intemacional de Agricultura Tropical (CIAT) 
Cali , Colombia (1989- presenl) 
Plant NutritionistlPhysiologist, Tropical Forages Program 
Responsible for identifying adaptive atfábules of tropical forage 
species to low fertility acid soils; developing reliable screening 
indices far plan! evaluaUon and plan! improvement; investigating 
plant-soil interre/ationships with raspee! fa recycling o( nutrients 
in pasture-based production systems. 
University of California, Berkeley, California, USA (1984-89) 
Assistant Specialist, Department of Plant and Soil Biology 
Research on mechanisms of plant adapta/ion to low nutrient 
supply in sugar beet and soybean; use of light scattering and 
chlorophyll fluorescence techniques for mineral deficiency 
diagnosis; role of phosphorous in photosynthesis, carbon 
partitioning and carbon exporl; limiting factors in photosynthesis. 
University of lIIinois 
Urbana-Champaign, IlI inois, USA (1982-83) 
Researeh Associate, Department of Plant Biol09Y 
Research on mechanisms of plant adaptation to water stress in 
sunflower; leaf nutrient status (particularly magnesium and 
potassium) and photosynthetic response to low /eafwater potentials. 
Cornell University, Ithaca , NewYork, USA (April-May, 1982) 
Boyee Thompson Institute tor Plant Researeh 
Visiting Scientist, Environmental Biol09Y 
Research on effects of sulfur dioxide (SO) pollution on 
photosynthesis, leaf conductance and slomatal metabolism of 
peas. 
ID 
Appendix .'1-1 
CJ~ - I.M Roo 
University of lIIinois at Chicago 
Chicago, IIlinois, USA (1981-82) 
Research Associate, Department of Biological Sciences 
Research on the role of light modulafíon of enzyrnes in the 
mechanisms of stoma/al movement; effecls of su/file and arsenife 
on sfomatal metabolism. 
International Crops Research Institute tor the Semi-Arid 
Tropics (ICRISAT) 
Patancheru, Andhra Pradesh, India (1979-81) 
Plant Physiologist, Pulse Physiology 
Research on developmenf of field screening fechniques lo 
evaluate pigeonpea genotypes for Iheir to/efanee to abiotic 
slresses, particu/arly sa/inify and waterlogging; growth analysis of 
pigeonpea hybn"ds and their parenls; response of pigeonpeas lo 
environmenf and cultural praetices such as spacing, plant 
population, irrigation and nutn"ents. 
Supervisory Experience : Supervision of Ph.D. (3) and Undergraduate degree (3) thesis 
projeets tor students regislered in the Universities from USA, 
Germany and Colombia. 
Areas of Specialization : Plant traits and meehanisms for adaptation lo low fertility acid 
soils; nutrient acquisition and nutrient cycling in pasture-based 
production systems; physiological responses of plants lo edaphic 
and climatic stresses. 
Professional Memberships: .. American Soeiety tor Plant Physiology 
.. American Sociely of Agronomy 
.. Crop Seienee Sociely of Ameriea 
.. Soil Seienee Society of Ameriea 
Education : Ph.D. Plant Physiology 
Sri Venkateswara University, India (1978) 
Dissertation Tifle: Carbon metabolism and water relations of six 
woody weeds and their modification by 
paraquat and 2,4,S-T. 
M.S. 80tany, Plant Physiology 
8hopal Universily, India (1973) 
8 .S. 80tany, Chemistry, Zoology 
Andhra University, India (1971 ) 
El 
Appendix A- I 
CV - J.M. Roo 
Languages: English -Fluent 
Telugu -Fluent 
Hindi -Conversational 
Spanish -Conversational 
Citizenship: India n 
Country of Residency : Colombia 
Publications: Rao, 1. M. and Kerridge, P.C. 1994. Mineral nutrition of forage 
(Author of 52 artlcles and 6 book Arachis. In: P. C. Kerridge and W. Hardy (eds.). The 
chapters, a list of last 5 years) biology and agronomy of forage Arachis. pp. 71-83. CIAT, 
Cal i, Colombia. 
Fisher, M.J., Rao, 1. M., Ayarza, M.A , Lascano, e. E., Sanz, J.!. , 
Thomas, R.T. and Vera, RR 1994. Carbon storage by 
introduced deep-rooted grasses in the south American 
savannas. Nature 371 : 236-238. 
Rao,1. M., Kerridge, P.C. and Macedo, M. 1994. Adaptation to low 
fertility acid soils and nutritional requirements of 
Brachiaria. In: The biology, agronomy, and improvement 
of Brachiaria . CIAT, Cali , Colombia, in press. 
Boddey, R.M., Rao, 1. M. and Thomas, R.J. 1994. Nutriant cycling 
and environmental impact of Brachiaria pastures. In: The 
biology, agronomy, and improvement of Brachiaria . CIAT, 
Cali, Colombia, in press. 
Rao, 1. M" Borrero, V., Ayarza, M.A and Garcia, R. 1994. 
Adaptation of tropical forage species to acid soils: The 
influence of varying phosphorus supply and soil type on 
plant growth. In: R. A Date, N. J . Grundon, G. E. Raymenl 
and M. E. Probert (Eds.), Plant-80illnteractions at Low 
pH: Principies and Management. Kluwer Academic 
Publishers, Dordrechl, The Nelherlands, in press. 
Rao, 1. M., Ayarza, M.A and Thomas, RJ. 1994. The use of 
carbon ¡solope ratios to evaluate legume contribution to 
soil enhancement in tropical pastures. Plant and 80i1162: 
177-182. 
Rao, 1. M. and Terry, N. 1994. Leaf phosphate status and 
photosynthesis in vivo: Changes in sugar phosphates, 
adenylates and nicotinamide nucleotides during 
photosynthetic introduction in sugar beet. Photosynthetica 
30: 243-254. 
ID 
Appendix A-J 
CII - J.M. Roo 
Rao, LM ., Zeigler, R.S., Vera , R. and Sarkarung , S. 1993 . Selection and breeding for acid-soil 
tolerance in crops: Upland rice and tropical forages as case studies. BioScience 43: 454-465. 
Rao, 1. M., Borrero, V., Ayarza, M.A and Garcia, R. 1993. Adaptation of tropical forage species to acid 
soils: The influence of varying phosphorus supply and soil type on phosphorus uptake and use. 
In: Barrow, N. J . (ed.) . Plant Nutrition - from Genetic Engineering to Field Practice. pp. 345-
348. Kluwer Academic Publishers, Dordrecht,The Nether1ands. 
Rao, I.M., Ayarza, MA, Thomas, R.J ., Fisher, M.J ., Lascano, C. and Borrero, V. 1993. Adaptation 
responses of tropical grass-Iegume associations to acid soits . Proc. XVlllntemational 
Grassland Congress, New Zealand & Queensland, Australia . pp. 1559-1560. 
Fisher, M.J. , Lascano, C.E., Thomas, R.J ., Ayarza , M.A , Rao, 1. M., Rippstein, G. and Thomley, H.J.M. 
1993. An integrated approach to understand soil-plant-animal interaetions on grazed legume­
based pastures on tropical aeid soils. Proc. XVII Intemational Grassland Congress, New 
Zealand & Oueensland, Australia . pp. 1903-1204. 
Thomas, RJ., Fisher, M. Lascano, C., Rao, I.M ., Ayarza, M.A. and Asakawa, N. 1993. Nutrient cyeling 
via forage litter in tropical grassllegume pastures. Proc. XVlllnternational Grassland 
Congress, New Zealand & Oueensland, Australia. pp. 508-509. 
Ayarza , M.A., Rao, I.M., Thomas, RJ ., Fisher, M.J., Lascano, C.E. and Herrera, P. 1993. Standing root 
biomass and root distribution in Brachiaria decumbens/Arachís pintoi pastures. Proe. XVII 
International Grassland Congress, New Zealand & Oueensland, Australia . p. 1921. 
Rao, LM. , Freeden, AL. and Terry, N. 1993. Influence of phosphorus nutrition on photosynthesis and 
carbon allocation and partitioning in sugar beet and soybean grown with a short photoperiod. 
Plant Physiology and Biochemistry 31 : 223-231 . 
Rao, LM., Ayarza, M.A., Thomas, RJ ., Fisher, M.J ., Sanz, J.L, Spain, J.M. and Lascano, C.E. 1992. 
Soil-plant factors and processes affecting produetivity in ley farming. In: Pastures for the 
tropical lowlands: CIAT's contribution. pp 145-175. CIAT, Cali, Colombia. 
Rao, I.M ., Roca. W .M. , Ayarza , M.A , Tabares, E. and García, R 1992. Somaclonal variation in plant 
adaptation to acid soil in the tropical forage legume Stylosanthes guianensis. Plant and Soil 
146: 21-30. 
Terry, N. and Rao, LM. 1991. Nutrients and photosynlhesis: iron and phosphorus as case studies. ln: 
J.R . Porter and D.W. Law10r (eds) , Plant Growth: Interactions with Nutrition and Environment. 
pp. 55-79. Cambridge University Press, Cambridge, England. 
Rao, LM ., Fredeen, AL. and Terry, N. 1990. Leaf phosphate status, photosynthesis and carbon 
partitioning in sugar beet 111. Diumal changes in carnon partitioning and exporto Plant Physiol. 
92:29-36. 
Rao, I.M ., Arulanantham, AR. and Terry, N. 1990. Diurnal changes in adenylates and nicotinamide 
nuc!eotides in sugar beet leaves. Photosyn. Res. 23:205-212. 
El 
Appendix ..1 - / 
O· - U. f. Roo 
Arulanantham, A.R. , Rao, I.M. and Terry, N. 1990. Limiting factors in photosynthesis VI . Regeneration 
01 ribulose 1, 5-bisphosphale limils pholosynlhesis al low pholoehemical capacity. Planl 
Physiol. 93:1466-1475. 
Fredeen, AL., Raab, T.K., Rao, I.M. and Terry, N. 1990. Elleels 01 phosphorus nulrilion on 
photosynthesis in Glycine max. Planta 181 :399-405. 
Rao, I.M., Arulanantham, AR. and Terry, N. 1989. Leaf phosphate status, photosynthesis and carbon 
partitioning in sugar beet 11. Diurnal changes in sugar phosphates, adenylates and nicotinamide 
nueleolides. Planl Physiol. 90:820-826. 
Fredeen, Al. , Rao, 10M. and Terry, N. 1989. Influence of phosphorus nutrition on gro'Wth and carbon 
partilioning in G/ycine max. Planl Physiol. 89:225-230. 
Rao, I.M ., and Terry, N. 1989. Leaf phosphate status, photosynthesis and carbon partitioning in sugar 
beet 1. Changes in growth, gas exchange and Calvin cycle enzymes. Plant Physiol. 
90:814-819. 
m 
Appendix A-2 c.v. 
c .... tro Intemoc:lc;tlo! de AQllc\lllufo TtopIcol 
~, ....~ c;tlgl C..,t. lo! Tropical ."""eulll,or. 
Horst J. Marschner 
Pesitlon in Project: German Project Coordinator 
International Research 
Experience: Member of the Deutsche Akademie der Naturforscher ~ 
Leopoldina 
1993 
University Hannover 
Dr. rer hort. honoris causa, Faculty of Horticulture. 1989 
University of Hohenheim 
Professor ef Plant Nutrition. Since 1977 
Commonwealth Scientific and Industrial Research 
Organization (CSIRO), Adelaide, Australia 
Division of Horticultural Research. 1973-1974 
Technical University af Berlín 
Institutie of Plant Nutrition. 1966·1976 
Full Professor and Director 
University of California, Berkely, USA 
Department of Soil Science and Plant Nutrition. 1965·1966 
Research fellow 
University of Hohenheim 
Institute of Plant Nutrition. 1960·1966 
Associate Professor 
German Academy of Sciences, Berl in 
Institute of Crop Research Gatersleben. 1958-1960 
Research Associate 
University of Jena 
Institute of Agricultural ehemistry. 1955·1958 
Research fellow 
m 
Appendix A-2 
el ' - Horst Marschner 
Education : University of Jena 
Study 01 Agriculture and Chemistry 
1951-1955 
Ph.D., University of Jena 
1957 
Main Research Interests: Mineral metabolism of plants (uptake, translocation, function of 
mineral elements). Genotypical differences in mineral 
metabolism of crop plants (deficiency, toxicity). Adaptatian af 
plants lo adverse soil canditians (acid ar alkaline sails, salinity, 
water lagging). Yield farmalian f 'saurce-sink-relatianship"). 
Environmental aspects af ferti lizer applicatian (nitragen, heavy 
metals). Plant-micrabial symbiasis: Endo- and 
ectomycorrhizae, Rhizobium. 
Joint research projecls on plant nutrition and crop production, 
Universities and Research Institulions, e.g. in the P.R. of China 
(increase in efficiency of fertilizer appllcation), Israel (i ran 
acquisition; drought stress tolerance) , Niger (increase in 
nutrient use efficiency), Syria (phosphorus efficiency), Turkey 
(micro-nutrient supply in citrus and coltan, zinc efficiency), and 
Colombia (phosphorus efficiency of pasture legumes and 
grasses). 
Languages: German - Fluent 
English - Fluent 
Citizenship: German 
Country of Residency: 
Germany 
ID 
Appendix A-} 
e l ' - Horst .\ lorsch"er 
Publications: 
301 publications in naliona1 and ¡ntemat¡onal scientific joumals. Textbook: Mineral Nutrition of Higher 
Plants, Academic Press, first edifion 1986; second edition 1995 (in press) . 
Relevant Publications 
119 publications forthe past 5 years 
Marschner, H. and ROmheld, V. Strategies o( plants foracquisition of ¡ron. Plant Soi1165, 261-
274 (1994) 
Marschner, H. and Dell , B. Nulrienl uplake in mycorrhizal symbiosis. Plant Soi1159, 89-102 (1994) 
Dinkelaker, 8 .. Hahn, G. and Marschner, H. Non-destructiva methods for demonstrating chemical 
changes in Ihe rhizosphere. 11. Application 01 methods. Plant SoiI 155/ 158, 71-74 (1993) 
Marschner, H. Nutrient dynamics al the soil·root interface (rhizosphere) . In: Mycrorrhizas in 
Ecosystems, D.J. Read el al. eds., pp. 3-12. CAB Intemational, Wallinglord Oxon, UK. 
(1992) 
Marschner, H. Mechanisms of adaplalion of planls lo acid soits. Plant SOill34, 1-20 (1991). 
Marschner, H. Nutrient dynamics in the rhizosphere. 3 Simposium Nacional sobre Nutrición 
Mineral de las Plantas, Palma (Baleares) 18.-20. Sepl. pp. 1-13 (1990). 
Marschner, H., ROmheld, V. and Zhang F.S. Mobilization of mineral nutrienls in the rhizosphere 
1 
by rool exudates. In: Ed. Masyoshi Koshino. Transactions 14TH Intern. Conference of 
Soil Science, Kyoto, Japan, Vol. 11 , pp. 158-163. (1990). 
I 
Ernst, M., Rómheld, V. and Marschner, H. Estimation o( phosphorus uptake capacity by differenl 
zones of Ihe primary rool of soil-grown maize (Zea mays L.) Z. Pflanzenemahr. Bodenk. 
I 152, 21-25 (1989) 
Dinkelaker, V., Romheld, M. and Marschner, H. Citric acid excretion and precipitalíon of calcíum 
I cítrate in the rhizosphere ofwhite tupín (Lupinus albus L.) Plant Cell Environ. 12, 285-
292 (1989) 
Marschner, H. and Cakmak, 1. High lighl intensity enhances chlorosis and necrosis in leaves of 
zinc, potassium and magenesium deficient bean (phaseolus vulgaris) plants. J. Plant 
Physiol. 134, 308-315 (1989) 
Marschner, H. and Haussling. Organic and inorganic soil phosphates and acíd phosphatase 
activity in the rhizosphere of 80-year-old Norway spruce (Picea abies (L» Karst.) trees. 
I Biol. Fertif. Soits 8, 128-133 (1989) 
I 
m 
.UNIVERSITAT HOHENHEIM 
INSTITUT FÜR PFLANZENERNAHRUNG 
Prof. Dr. Dr. h. c.. Horn Mar~chner 
Appendix B 
Dr. P. Kerridge Partner Conflrmation Letter 
Leader of the Tropical Fo rages Program 
Fruwin.hnr. 20 
C 1 A T 70599 SWttpn. 
A . A. 6713 
Cal!, COLOMBIA 
11.1.1995 
Oear Dr. Kerridge, 
I I I am writing to eonfi~ my c ommitme nt and that of the 
Institute of Plant Nutrition of this Univer sity, to the 
special project entitled 
"Calcium acquisition and utilizatio n by forage 
grasses and legumes" 
whieh will be proposed to DMZ. 
Yours sincerely, 
~ 
1 
El 
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