Plant and Soil 202: 133–147, 1998.
© 1998 Kluwer Academic Publishers. Printed in the Netherlands.
133
Leaf area expansion and assimilate production in sunflower ( Helianthus
annuus L.) growing under low phosphorus conditions
Daniel Rodr´ ıguez
1,4
, M.M. Zubillaga
1
, E.L. Ploschuk
2
, W.G. Keltjens
3
, J. Goudriaan
4
and R.S.
Lavado
1
1
Fertilidad y Fertilizantes, Facultad de Agronom´ ıa U.B.A. Av. San Martin 4453, Buenos Aires (1417), Argentina,
2
I.F.E.V.A., Facultad de Agronom´ ıa U.B.A.,
3
Sub-department of Soil Science and Plant Nutrition, Wageningen
Agricultural University, Dreijenplein 10, 6703 HB, Wageningen, and
4
Sub-department of Theoretical Production
Ecology, Wageningen Agricultural University, P.O. Box 430, 6700 AK, Wageningen, The Netherlands
∗
Received 30 June 1997. Accepted in revised form 24 April 1998
Key words: leaf expansion rate, modelling, phosphorus, photosynthesis, sunflower
Abstract
Reductions in leaf area and plant growth as a consequence of phosphorus (P) limitations have been attributed both
to direct effects of P shortage on leaf expansion rate and to a reduced production of assimilates required for growth.
Canopy assimilation and leaf area expansion are closely interrelated processes. In this work we used experimental
and simulation techniques to identify and study their importance in determining leaf area on sunflower ( Helianthus
annuus L.) growing under P-deficient conditions. Experiment 1 was done outdoors, in Buenos Aires, Argentina,
and Experiment 2 in a glasshouse in Wageningen, The Netherlands. In both experiments we studied the effects of
soil P addition on leaf appearance, leaf expansion, dry matter accumulation, and leaf photosynthesis of non-water
stressed plants grown in pots containing a P-deficient soil. Before sowing the equivalent amounts of 0–600 kg of
super phosphate ha
−1
were added to the pots. Phosphorus deficiency delayed leaf appearance increasing the value
of the phyllochron (PHY) up to 76%, the rate of leaf area expansion during the quasi-linear phase of leaf expansion
(LER) was reduced by up to 74%, with respect to high P plants. Phosphorus deficiency reduced by up to 50% the
rate of light saturated photosynthesis per unit of leaf area (AMAX) in recently expanded leaves, while at low
levels of leaf insertion in the canopy, AMAX was reduced by up to 85%, when compared to that in high P plants.
Phosphorus deficiency also reduced the duration of the quasi-linear phase of leaf expansion by up to eight days.
The values of LER were related (r = 0.56, P < 0.05) to the mean concentration of P in all the leaves (Leaves P%)
and not to the concentration of P in the individual leaf where LER was determined (r = 0.22, P < 0.4) suggesting
that under P deficiency individual leaf expansion was not likely to be regulated by the total P concentration at leaf
level. The values of AMAX of individual leaves were related (r = 0.79, P < 0.01) to the concentration of total P in
the corresponding leaf (Leaf P%). LER showed a hyperbolic relationship with Leaves P% (R
2
= 0.94, P < 0.01,
n = 13) that saturate at 0.14%. AMAX showed a hyperbolic relationship with Leaf P% (R
2
= 0.73, P < 0.01, n =
53) that saturated with values of Leaf P% higher than 0.22. A morphogenetic model of leaf area development and
growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and
to study the effects of model parameters on the growth of sunflower plants under P-deficient conditions. With this
model we identified the existence of direct effects of P deficiency on individual leaf area expansion. However, we
calculated that under mild P stress conditions up to 83% of the reduction in the observed leaf area was explained
by the particular effects of P% on the rate of leaf appearance, on the duration of the linear period of leaf expansion,
∗
Present address of corresponding author. FAX No: 31 317
484892. E-mail: Daniel.Rodriguez@staff.tpe.wau.nl