Plant and Soil 205: 155–162, 1998.
© 1998 Kluwer Academic Publishers. Printed in the Netherlands.
155
Phosphorus supply and the growth of frequently defoliated white clover
(Trifolium repens L.) in dry soil
Dhananjay K. Singh
1,2
and Peter W. G. Sale
1
1
Department of Agricultural Sciences, La Trobe University, Bundoora, Victoria 3083, Australia and
2
Frank Wise
Institute, Kununurra, Western Australia 6743, Australia
Received 29 July 1997. Accepted in revised form 7 September 1998
Key words: defoliation frequency, phosphorus deficiency, soil water stress, water and P uptake, white clover growth
Abstract
A previous study found that increased phosphorus (P) supply to frequently defoliated white clover plants, growing
in a low-P, dry soil, alleviated water stress symptoms and increased plant recovery on rewatering. In this study we
determined how these stresses influence white clover growth. Measurements were made of the leaf canopy, stolon
infrastructure and root system of the white clover plants growing in a low-P soil. Treatments included the factorial
combination of four levels of P supply, two defoliation frequencies and two soil water treatments. White clover
growth declined markedly when P-deficient plants were exposed to frequent defoliation and dry soil conditions.
Leaf area was more affected than other parameters, in that the combination of stresses reduced leaf area to 2%
of maximum observed for infrequently defoliated plants growing in high-P soil, with adequate water. Increased P
supply generally increased the growth of all plant parts. Frequently defoliated plants growing in dry soil produced
similar or greater leaf mass and leaf area as plants from similar treatments growing in wet soil, when the P supply
increased to 50 mg P kg
−1
soil. Higher P rates were able to negate the effect of dry soil on these frequently
defoliated plants, as a result of larger water and P uptake. Also, the frequently defoliated plants with restricted root
growth did not respond to a small increase in P supply (17 mg P kg
−1
soil) for the leaf growth, irrespective of
whether they were growing in wet or dry soil. Infrequently defoliated plants with greater root growth, compared to
frequently defoliated plants, more than doubled their leaf mass with this P treatment.
Introduction
Even in the higher rainfall regions of south east Aus-
tralia, it is not uncommon for pastures to be subjected
to dry soil conditions. The restriction in forage supply
during these periods can result in frequent grazing by
livestock. Recent findings suggest that the persistence
of white clover (Trifolium repens) under these condi-
tions can be improved by increasing P supply to the
plants. For example, Singh et al. (1997) found that fre-
quently defoliated, low-P plants in dry soil displayed
severe water stress symptoms, and many plants died
following the desiccation and senescence of most of
the leaves. On the other hand, high-P plants not only
survived and recovered well on rewatering, but they
∗
FAX No: 3 9471 0224. E-mail: agrds@lube.latrobe.edu.au
were also better able to maintain a higher leaf water
potential and extract more water from a drier soil.
Increased P supply was found to maintain shoot
growth of white clover under frequent defoliation
regimes, although overall shoot growth was reduced
by frequent defoliation (Caradus and Snaydon, 1988;
Singh and Sale, 1997). The higher P supply resulted in
increased numbers of smaller leaves and an increase
in stolon branching in these plants (Singh and Sale,
1997). Other studies show that an increased P sup-
ply improves clover growth in dry soil (Simpson and
Pinkerton, 1989 for subterranean clover; Dodd et al.,
1992 for white clover). However, the effect of P sup-
ply on the growth of white clover that is subjected to
both frequent defoliation and dry soil conditions to-
gether has not been studied. The question is whether
increased P supply can negate the effect of dry soil on
the plant growth, given that a high-P supply alleviated