Environmental control of canopy dynamics and photosynthetic rate in the evergreen tussock grass Stipa tenacissima

Seasonal changes in leaf demography and gas exchange physiology in the tall evergreen tussock grass Stipa tenacissima, one of the few dominant plant species in the driest vegetation of Europe, were monitored over a period of two years at a field site in semi-arid south-eastern Spain. Three age-classes of leaves - young, mature and senescent - were distinguished in the green canopy. Production of new leaves and extension growth of older leaves occurred exclusively from October-November to May-June. The rate of extension was significantly correlated with gravimetric soil water content. Leaf growth ceased after gravimetric soil water content fell below 0.015 g g(-1) at the beginning of the dry season which corresponded to pre-dawn leaf water potentials of -3.0 MPa. Leaf senescence and desiccation reduced green leaf area by 43-49% during the dry season. Diurnal changes in the net photosynthetic rate of all three cohorts of leaves were bimodal with an early morning maximum, a pronounced midday depression and a small recovery late in the afternoon. Maximum photosynthetic rates of 10-16 micromol CO(2) m(-2) s(-1) were attained from November 1993 to early May 1994 in young and mature leaves. Photosynthetic rate declined strongly during the dry season and was at or below compensation in September 1994. Gas exchange variables of young and mature leaves were not significantly different, but photosynthetic rate and diffusive conductance to water vapour of senescing leaves were significantly lower than in the two younger cohorts. Leaf nitrogen content of mature leaves varied seasonally between 2.9 and 5.2 g m(-2) (based on projected area of folded leaves), but was poorly correlated with maxima of the photosynthetic rate. There was a stronger linear relationship between the daily maxima of leaf conductance and pre-dawn leaf water potential than with atmospheric water vapour saturation deficit. Seasonal and between-year variation in daily carbon assimilation were caused mainly by differences in climatic conditions and canopy size whereas the effect of age structure of canopies was negligible. Since water is the most important limiting factor for growth and reproduction of S. tenacissima, any future rise in mean temperature, which might increase evapotranspiration, or decrease in rainfall, may considerably reduce the productivity of the grasslands, particularly at the drier end of their geographical distribution.