The seasonal regulation of gas exchange and water relations of field grown macadamia

Macadamia F. Muell is a recently domesticated nut crop characterized by a certain amount of drought tolerance. Whilst stomatal regulation of gas exchange and water relations has been documented in macadamia, there are no reports on non-stomatal limitations to photosynthesis and if changes occur over a season in relation to different phenological stages. It was hypothesised that in order for macadamias to be adapted to a seasonally dry native environment the trees would be characterised by strong stomatal control over gas exchange, indicating an isohydric water use strategy that is related to a hydraulic limtation within the tree. However, due to high assimilate demand during nut filling, the level of stomatal control would vary between fruiting and non-fruiting phenological stages. Gas exchange and water relations measurements were made for 18 months on irrigated mature macadamia trees (cv. HAES 695, 'Beaumont') in a subtropical region of South Africa. Results confirmed that macadamias had relatively low light-saturated net CO2 assimilation rates (A(max)) (8.34 +/- 1.21 mu mol CO2 m(-2) s(-1)). The low A(max) values resulted from relatively high stomatal and non-stomatal limitations and decreasing stomatal conductance (g(s)) in response to increasing leaf-to-air vapour pressure deficit (VPDleaf ) above ca. 2 kPa. Strict stomatal control and nearly constant midday leaf water potential (psi(leaf) = -1.16 +/- 0.43 MPa) confirmed the predominantly isohydric nature of the crop, which seemed to be a result of low hydraulic conductance in the stem to leaf pathway. Significant differences in leaf gas exchange capacity were, however, observed between the fruiting and non-fruiting phenological stages. The presence of fruit resulted in significantly higher A(max) (10.27 +/- 2.23 mu mol CO2 m(-2) s(-1)) compared to non-fruiting periods (A(max) = 6.58 +/- 2.00 mu mol CO2 m(-2) s(-1)). Increases in A(max) were mediated by increased rates of electron transport (J(max)) and triose phosphate use (TPU). Fruiting stages were also characterized by variable responses of g(s) to increases in VPDleaf, which indicated varying degrees of isohydricity. This study reaffirms that macadamias are inherently adapted to seasonally dry environments, characterized by strict stomatal control, yet under humid conditions or in the presence of developing fruit, macadamias are proposed to deviate from a purely isohydric water management strategy.