Nine lines of peanut (Arachis hypogaea L.) including four high yielders, one non-nodulating line and its progenitor parents, one high nitrogen fixer and a national check, were studied in situ for 14c translocation to various plant's parts- leaves, stem, root nodule, shell and kernel - at peg development and harvest stages. Of the 68% of observed yield variation accounted for by the carbon translocation to roots, nodules, shells and kernels at harvest, 65% was accounted by the former two traits alone. The relative increase in translocation to roots, nodules, shells and kernels at harvest, 65% was accounted by the former two traits alone. The relative increase in translocation to roots and nodules at harvest over peg development stage directly influenced pod yields. This was substantiated by the nature and magnitude of correlations between pod yield (PY) and % 14c at peg development (PD) and harvest (HS) stages. There was no correlation between PY and % 14c in (root + nodule) at PD; but that correlation at HS was positive and significant. The observed differences in 14c partitioning between the high and low yielders suggest partitioning of carbon to reproductive parts as an additional economic selection criterion for improving productivity in peanuts

Arunachalam, V.

Vinod Prabhu, K.

English

Publications of the IAS Fellows

~r----.STER NEGATIVE BER: 09296.16 
Arunachalam, V. and Vinod Prabhu 
Journal 0 Oilseeds Research, 8 1992: 
199-204. 
Record no. D-80 
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J. Oilseeds Res 8(1991) 199-204 
ISSN 0970 .. 2776 
CARBON.· TRANSLOCATION ACCOUNTING FOR YIELD 
V ARIATI~)l~·IN PEANUT (ARACHIS HYPOGAEA L.) 
. . 
K. VNOD PRABHul and V. ARUNACHALAM· 
, ., .. 
Division of GeneticS~ Indian Agricultural Research Institute, New Delhi 110 012. (India) 
. . 
.. 
ABSTRACT 
Nine lines. of peanut (Arachis hypogaea L.) including four high yielders, one noli-
nodulatin~ line and its progenitor parents, one high nitrogen fixer and a national check, 
were studied in situ for 14c translocation to various plant's parts-Ieaves,stem, root nodule, 
shell and kernel - at peg development and harvest stages. .Of the 68 % of observed yield 
variation accounted for by the carbon translocation to roots, nodules, shells and .kernels 
at harvest, 65 % was accounted by the former two traits alone. The relative increase in 
translocation to roots and nodules at harvest over peg dev~lopment stage directly influ-
encedpod yields. This was substantiated by the nature and magnitude orcorreIations 
between pod yield (PY) and % 14c at peg development (PD) and harvest (HS) stages. 
There was no correlation between PY and % 14c in (root+nodule) at PD; but that 
correlation at HS was positive and significa.nt. The observed differences in 14c . 
partitioning between the high and low yielders suggest partitioning. of carbon to 
reproductive parts as an additional economic selection criterion for improving productivity 
in peanuts. . 
Key words Peanut; Yield potential; Selection; Assimilate partitioning; Carbon trans-
location. 
INTRODUCTION 
Amopg ~he factor& determining pod yield in peanut, partitioning of daily assi .. 
milates to the growing fruit is of crucial importance. This inference was arrived at 
from a phYsioI.ogical analysis of yield differences of popular varieties released in the 
U.S.A. in the distant and recent past (Duncan et al., 1978). While Dixie runner, one 
of the earliest varieties with an yield potential of 2.4 t/ha had a partioning factor of 
41 %, Early ~unch, a variety released in the recent past with an yield potential of 5.4 
tjha had a very high partitioning factor of 99 %. Even under conditions limiting net 
photosynthetic rate, it has been found possible to increase inter-organ partitioning 
of dry matter "from unusable vegetative· parts to commercially useful sinks {Gifford 
et al., 1984). Experimental evidence was wide"to support this view (Graham, 1"978; 
Carlson and Brun, 1984; Crafts-Bradner e{ al. , 1984). It will then be feasible to use 
translocation of assimilateS to growing fruit as a selection criterion to improve producti-
vity in peanuts. 
. . 
At the Indian Agricultural Research Institute, a number of high yielding peanut 
lines has been derived in the research programme of a national project on peanuts. 
Four of those lines and a few genetically diverse lines dIffering in yield.were used as 
the test material to ascertain the extent to which carbon translocation could account for 
the . observed yield differences and wheather it could be a useful selection criterion. The 
. results a~ reported here. 
• J 
1 
e 
1 Present address: IARI Regional Station, Tutikandi, Simla 171.004, India. 
Received on 6.12.1991 
200 Journal of Oilseeds Research 
MATERIALS AND METHODS 
Nine lines' of peanut consisting of four developed in a national project (NFG 7, 
NFG D, NFP 140, 1441-A-l), a non-nodulating line, Non-nod and its progenitor 
parents,Nc 17 and PI 259747, a national check, Robuf33-1 and a high nitrogen fixing 
accession from North Carolina, NC Ac 2821 were grown in a randomised block design 
with three replications. The crop received a basal dressing of 50 kg P20S and 30kg 
N (as the available N in the soil was very 'low at 129 kg/ha) as per the national reco-
mmendations. At pegging stage, 250 kg gypsum (Ca S04) was added. The crop was 
grown after the onset of monsoon. under recommended cultural practices. The 
lines were sown in single row plots of 5m length on ridges spaced 60cm and plants 
within a ridge spaced 10cm apart. 
All the lines initiated flowering by 40 days after sowing (DAS). Pegs started 
developing 90 DAS and the crop reached harvest stage around 120 DAS. The peg 
development (90 DAS; PD) and harvest (120 DAS; HS) stages were us~d for measuring 
carbon translocation to plant parts. T 
In situ 1'4c isotope feeding was employed to measure translocation. The iso-
tope was fed on two plants per plot in the field (which was irrigated two days before 
sampling) after caging the plants in special polythene chambers measuring 30cnl x 
.... 30cm. X 30cm. Isotope solution in the form of sodium carbonate (28. 25m ci/m mole) 
was diluted to give 30(1. ci/ml. 200mg non-radioactive sodium carbonate and 4mI disti-
lled water were added l:ml isotope solution. 14C02 was released by mixing 8m! of 70 % 
perchloric acid. The whole process had earlier been standardised by the radiotracer 
unit of the Department of Agricultural Chemistry, Andhra Pradesh Agricultural Uni-
versity, Hyderabad, India. 
Each sample was incubated for 40 minutes. 24 hours after isotope feeding, the 
plants were pulled out and separated into leaves (LF), stem (SM), roots (RT), nodules 
(ND), shell (SH) and kernel (KL). The samples were dried in a hot air oven. The 
dried plant parts \\-ere separately ground fine and one g dry samples were exposed 
• 
under a Geiger-Muller Counter l GCS 13) for 200 seconds at 1410 working volts. 
The distribution of 14C was estimated taking into account the dry weight of 
plant parts. For example, Zi J the % 14C in plant part i was calculated as: 
z·= 1 
w·x. 1 1 x 100 
where Wi was the dry weight and Xi the isotope count of plant part i, i=1,6. i=l 
refers to LF, 2 to SM, 3 to RT, 4 to ND, 5 to SH and 6 to KL. 
Dry pod yields of the sampled plants were also noted. _ 
Vinod Prabhu and Arunachalam 201 
RESULTS AND DISCUSSION 
There was considerable variation among the 9 lines for dry weight and carbon 
translocation to plant parts as reflected in their mean values (Table 1). The lines, NFG 
7, NFP D, 1441-A-1, Robut 33-1 and NC Ac 2821 gave pod yields per plant above the 
experimental'mean (30.4 g plant-I). In general about 75 % of 14c was translocated to 
leaf and stem with individual variation among lines. More than 80 % of 14c were found 
in leaf and stem in the lines NC 17, Non-nod and NFP 140. High % 14c in the plant 
organs, root, nodule, shell and kernel, was observed in NFG 7 (31.3 %) and Robut 
33·1 (25.4%). 
TABLE 1 Per .cent dry weight (D) and 14c (T) in different plant parts at harvest in peanut. 
Line 
NC Ac 2821 
NFG 7 
NFPD 
NFP 140 
1441-A·1 
NC 17 
PI 259747 
Non-nod 
Robut 33-1 
Mean 
SE ± 
-
LF + ST RT + ND SH + KL Pod yield 
-- ---- --- -------- (gjplant) 
D T D T D T 
------- --- ---
50.4 77.2 2.5 4.6 47.1 18.2 31. 9 
51.2 68.7 2.4 6.0 46.4 25.3 45.3 
51 . I 76.5 2.4 4.7 46.5 18.8 36.2 
53.2 80.5 3.0 3.2 43.8 16.3 24.0 
53.5 67.2 3.0 3.8 43.5 21.0 36.1 
58.4 83.8 3.1 4.1 38.5 12.1 21.4 
60.1 75.3 4.3 2.8 35.6 21.9 19.9 
67.9 80.9 3.6 2.1 28.5 17.0 9.5 
50.3 74.6 2.5 4.3 47.2 21.1 49.2 
---------------_ .. ---.--------------------
55.1 
5.94 
76.1 
5.48 
3.0 
0.64 
4.0 
1.15 
41.9 
6.46 
19. I 
3.81 
30.4 
8.04 
-_._---------_ .. --_._---------
Dry weights were distributed among above ground (leaf+sten1) and under-
ground (root+nodule+shell+kerneI) organs almost equally (Table 1). Non-nod 
provided an exception where the dry weight of leaf and stem was about double that 
of the rest of the organs. 
High yielding lines like NFG 7 and Robut 33.11 partitioned relatively high carbon 
to root, nodule l shell and kernel while the opposite was true of low yielders like NC 
17 and Non-nod. In some lines such a direct association was not apparent. For 
example, carbon patritioned to underground parts was 24.8 % for 1441-A-l with an 
yield of 36.1 g plant- t • This difference for PI 259747 with an yield of 19.9 g plant-to 
This difference was complemented by a higher dry matter accumulation in 1441-A-l 
in pod (SH+KL), double that of PI 259747 at harvest. As inferred in grain legumes 
202 Journal of Oilseeds Research 
like chickpea and pigeonpea, efficient mobilisation of carbon and nitrogen froID. stem 
and leaves and high photosynthetic and N-fixing rates during seed filling 'would also 
be the other important factors for explaining differences in productiv,ity (Singh, 1990). 
Maximum C and N fixation occurred during flowering and early fruiting phases in 
. , 
chickpea while about half of the net photosynthate and N Were fix~d in t,he vegetative 
. . , . 
period itself in pigeonpea allowing for better remobilisation to pods at periods of seed 
fill explaining its comparatively higher yields. 
The association between pod yield and 14c and in various plant parts varied 
between peg development and harvest stages (1 able 2). While none of the correlation 
coefficients between pod yield and 14c in (leaf+stem), ,(root+nodule) or (shell+ 
kernel) was significant at peg development stage, the former two was significant at har-
vest. In particular, there was a positive and significant correlation between 14c in 
(root+nodule) and pod yield at harvest. 
TABLE 2. Correlation coefficients (r) between % 14c in various plant parts and pod yield at peg 
development (PD) and harvest (HS) stages in peanut. 
----------------------------------------------------------------
r PD HS 
-----------------------------------------------------------------
PY, C in (LF + ST) 
PY, C in (RT + ND) 
PY, C in (SH + KL) 
*Significant at P=0.05 
-0.55 
o 
0.52 
-0.67* 
0.81* 
0.60 
Such an association was lacking between dry weight of plant parts and 14c in 
them at both stages {Table 3). The only exception was a negatiVe correlation between 
dry wt::ight and 14c of (leaf+stem) at harvesL There was thus no premise a priori on 
which dry weight of plant parts could substitue 14c in them as selection criteria. 
TABLE 3. Correlation coefficients (r) between dry weights and % 14c in various plant 
parts at peg development (PD) and harvest (HS) stages in peanut. 
---
r 
LF + ST 
RT+ND 
. SH + KL . 
PD 
-0.63 
o 
0.64 
HS 
-0.75* 
0.48 
0.47 
. 
-------------------------'----,--------
*Significant at P=O.05 
, . . 
. The relative efficiency of several linear regression equations in accounting for the 
observed variation in pod yield was then compared. Only two equations, numbered 
11 and 3 (Table 4) .. could account, with a single independent variable,. for more than 
Vinod Prabhu and Arunachalam 203 
50 % of vatiation in pod )ield. Of these two, the linear regression equation between 
pod yield and % 14c in (root+nodule) at harvest was the best accounting for 65.5 % of 
variation in .pod yield. When 14c in (shoot+kernel) was also included the coefficient 
of determination could rise only to 68.4 % not justifying their inclusion despite the extra 
cost and time in measuring them. But the relative increase in 14c in (root+nodule) 
at harvest over peg development stage was found to be an important factor in deter .. 
mining pod yield (R2=54 %1 Table 4). 
TABLE 4. Regression equations of pod yield (Y) on percent 14c (x) partitioned to roots and nodules 
in peanut. 
------------ ------_._------
Regression equation R2 
---- -----------
(~j xl 
x2 
x3 
R2 
1 Y = - 5 . 2 + 8. 98 * X 1 ~!/ 65.5 
68.4 
54.0 
2 Y = -38.6 + 1.61* X2'!-i 
3 Y = 21. 2 + O. 1 7 * x 3 ~, 
. 
. 
--~-----.--------------- --------
14c in (root + nodule) at harvest; 
14c in (root + nodule + shell + kernel) at harvest; 
Relative increase in 14c in (root+nodule) at harvest over peg development stage. 
Coefficient of determination 
The results led to two major inferences: 
a. Pod yield was a function of partitioned carbon to roots and nodules. 
b. The increase in partitioning to roots and nodules from peg development 
to harvest stages had a substantial influence on realised pod yield~. 
Evidence to support partitioning of assinlilates to reproductive parts as a se-
lection criterion is abundant in published literature. It& use in constructing productive 
legunle plant types had earlier been recognised (Grahanl, 1978). Where efforts were 
unsucce&sful at increasing the rate of photosynthesis, inter-organ partitioning of photo-
synthates was used as a successful nlethod to realise conlmercial sinks (Gifford et a1., 
1984). Higher rate of photosynthate translocation to developing pods during peak 
filling period (corresponding to pod development stage of this study) resulted in higher 
yields of groundnut mutants compared to other derived genotypes (Lodha et al., 1983). 
In generat at the pod filling sta ge 1 a majority of 14c is required to be translocated to pods 
as plant parts compete for carbon among themselves for assimilation into specific sinks 
(Russell and JohnsoD, 1975). When accumulated carbohydrates in the stem Were not 
remobilised for seed development, poor yield resulted in Phaseolus (Adams et al., 1978). 
Selection of genotypes with more specific sink demands (where leaves translocated most 
photosynthates to pods) has been practised in soybean (Carlson et a/. J 1984) and cow-
pea (Pate et al., 1984). Published evidence suggests the need to select genotypes with 
,. 
204 Journal of Oilseeds Research 
profuse pod formation to generate more sink demand as was done in soybean (Carlson 
et al.,.1984J o~ that the sink demand itself did not exhaust the available source resulting 
in theret.ention of photosynthates in the leaves a~ late pod filling perio4 (Egli et al., 1980) 
.which limits realisation of the full geno~ypic potential. . 
Selection that can be operated before digging of pods saves a lot of time and 
labour in peanut research. In an earlier study (Prabhu et al., 1990) the utility of. leaf 
area and total nitrogen % measured 30 days after flOWering in selecting productive 
genotypes was highlighted. This investigation adds one more vital selection parameter 
namely, the rate and extent of carbon translocation to reproductive sinks around 
harvest. 
ACKNOWLEDGEMENT 
Our sincere thanks are due to Drs. K.V. Sathe and Badrinarayana Rao of the 
Radiotracer Laboratory, Andhra Pradesh Agricultural Universi~y, Hyderaba.d, India 
for their help in the in situ translocation studies. 
LITERATURE CITED 
ADAMS, M.W., WIERSMA, J.V. and SALAZAR, J. 1978. Differences in starch accumulation among 
dry bean cultivars. Crop Sci., 18 : 155-157. 
CARLSON,· D.R. and BRUN, W.A. 1984. Alteration of 14c assimilate paritioning in leaves of 
soybeans having increased reproductive loads at one node. Plant Physiol., 75 : 887-890. 
CRAFTS-BRANDNER, S.J., BELOW, F.E., HARPER, J.E. and HAGEMAN, R.H. 1984. Effect of 
nodulation on assimilate remobilisation in soybean. Plant Physiol., 76 : 452-455. 
DUNCAN, W.O, Mc CLOUD, D.E., McGRAW, R.L. and BOOTE, K.J. 1978. Physiological aspe-
cts of peanut yield improvement. Crop Sci., 18 ; 1015-1020. 
EOLI, D.B., LEGETT, J~E. and CHENIAE, A. 1980. Carbohydrate levels in soybean leaves during 
reproductive growth. Crop Sci., 20 : 468-473 . , 
GIFFORD, R.M. THORNE J.H., HITZ, W.D. and GIAQUINTA, R.T., 1984. Crop productivity 
and photoassirnilate partitioning. Science 205 : 801-808. . 
GRAHAM, P.H. 1978. Some problems and prospects of field beans (Phaseo/us vulgaris L.) in Latin 
America. Fied Crops Res., ] : 295 - 317. ~ 
. . 
LODHA, M.L., SHARMA, N.D, JOHARI, R.P., MEHTA, S.L. and NAIK, M.S. 1983. Nitrogen 
fixation and hydrogen mobilisation in the developing nodules of groundnut mutants. In Proc. 
National Symp. on Bioi. Nitrogen. Fixation, Indian Agricultural Research· Institute, New Delhi. 
pp. 216-228. ",' . " . 
PATE, J.S., ATKINS, C.A., LAYZELL, D.B. and SHELP, B.J. 1984. Effects of N2 deficiency 'on 
, . .transport and partitioning of C and N in a nodulated legume. Plant Physioi., 76:59-64. . 
..' , . 
PRABHU, K.V., ARUNACHALA·M, V. and BANDYOPADHYAY, A. 1990 .. Non-parametric 
. approach to ~ulti-trait seJectiol) for yield in groundnut (Arachsis hypogaea L.). Theor. Appl.~ 
Genet.: 80': 223-227. . . , ..' . 
, 
-. 


Carbon translocation accounting for yield variation in peanut (Arachis hypogaea L)

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Carbon translocation accounting for yield variation in peanut (Arachis hypogaea L)

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