Stomatal response to humidity: implications for transpiration

Abstract. Transpiration rates from apple leaves are analysed in terms of the ratio of latent heat flux (λ E ) to leaf net radiation ( Q ₁) and the climatological resistance ( r_i ). Increases in stomatal resistance with increasing leaf to air vapour pressure gradient ( D ), described by an empirical model, are incorporated in the analysis. This humidity effect causes the proportion of energy dissipated as latent heat to fall as Q ₁ increases, so that leaf transpiration rates in high energy environments are likely to be similar to those in lower energy environments. Boundary layer resistance ( r _a) exerts an increasingly important effect on transpiration rates as Q ₁ increases. At constant Q ₁ stomatal closure in response to increasing D results in very small changes in leaf temperature ( T ₁) across a wide range of ambient vapour pressure deficits (δ e ); r _a is then the major factor determining T ₁. The implications of these results are discussed.     

Water stress,leaf nutrients and defoliation: a model of dieback of rural eucalypts

Water stress, concentrations of foliar nutrients and damage of foliage by insects were studied over an eleven month period, for eight dieback eucalypts and eight closely matched healthy trees growing in close association on two grazing properties near Brisbane. Four healthy eucalypts in a neighbouring State Forest were also studied. The study region had suffered severe climatic stress between 1972 and 1976 when dieback was first observed by residents and when high populations of defoliating insects were observed. Dieback trees had a higher proportion of foliage damaged by insects and higher concentrations of foliar nutrients than did matched healthy trees. The study period was one of low climatic stress during which dieback and healthy trees followed similar seasonal patterns of predawn xylem pressure potential. Dieback trees developed lower daytime minimum xylem pressure potentials than did matched healthy trees, and differed in their stomatal responses. A model of initiation and development of rural eucalypt dieback is proposed. This may have general application to many non-specific rural diebacks in which heavy insect damage is implicated. The balance between rural eucalypts and their insect herbivores is precarious. Any factor capable of causing extensive defoliation, or an increase in foliar nitrogen, or an increase in populations of insect herbivores may upset this balance. A positive feedback loop may be activated, whereby the production of nitrogen rich epicormic foliage enhances a build up of insect populations. Repeated insect defoliation leads to tree dieback. Evidence in the literature supporting the model is reviewed and aspects requiring further research are outlined.     

Water Movement Through Plant Roots

Mathematical analysis of the hydraulics of water movement throughplant roots, in terms of radial and axial resistances, has ledto equations which provide new insights into the effects ofthe component resistan ces on water uptake by and movement throughindividual roots and root systems. The ratio of axial to radialresistance determines the optimum length of a root and its totalresistance to water movement. The equations permit direct calculationsof the plant water potentials necessary, at the base of theplant, to sustain given flow rates through root systems withgiven characteristics. Lateral spacing and the resistance ofindividual laterals are the dominant factors determining totalflux per unit area into a root. When soil water potential increases with depth (surface layersdrier) root resistance tends to decrease with increasing flowrate; the reverse occurs when the surface is wetter than thelower layers. Calculated patterns of water movement into andthrough roots, in relation to soil water potential and flowrate through the root, indicate efflux from root to soil undercertain conditions. This is considered to reflect reality, althoughthe fluxes are probably transient or intermittent. The equations presented should be combined with equations describingwater movement through soil to define the behaviour of the wholeroot-soil system adequately.     

Drought and dieback of rural eucalypts

The possibility that drought causes dieback of eucalypts in rural Australia was investigated. Water potential and canopy condition in dieback and healthy rural Eucalyptus blakelyi and E. melliodora trees were compared during and after an extreme drought in the ACT. All the trees were drought affected, but the extent was independent of the condition of their canopies at the beginning of the study.     

Regional and local variation in insect herbivory,vegetation and soils of eucalypt associations in contrasted landscape positions along a climatic gradient

Abstract Relationships with climate and local resources are developed for soils, vegetation and tree foliage as well as levels of herbivory for the dominant eucalypts at sites representing a regional gradient in climate and local contrasts in landscape position. Indicators of site productivity such as soil nitrogen and phosphorus, canopy height and cover, foliar nitrogen and water, and average leaf area tended to increase as climate became more favourable. Many were also higher in locally richer parts of the landscape. In contrast, specific leaf weight, an indication of sclerophylly, decreased as climate and local resources became more favourable. Rates of herbivory tended to increase with increasing site productivity and the associated changes in soil, vegetation and foliar properties, in broad agreement with models relating herbivory to resource availability and plant vigour. We found no evidence to support models relating high herbivory to low-resource environments and plant stress. The apparent level of herbivore damage on mature leaves was highest at intermediate levels of resources; this could reflect interactions between resource availability, rates of herbivory and rates of leaf replacement. Implications of these findings are discussed with respect to ways of measuring herbivory, regional patterns in rates and levels of herbivory, and the regional distribution of rural dieback associated with high herbivory.     

The Mechanisms of Apple Bud Morphogenesis: Analysis and a Model

Information in the literature suggests that the process of applebud morphogenesis is controlled by the level of growth substances,notably gibberellins in the bud, and by the availability ofgrowth substrate (S). The spur buds of apples must contain acertain minimum number of nodes (N_m) before flower primordiacan be formed, and the process of bud morphogenesis can be describedby a rate of node production with time, t by the equation It is postulated that the parameter k_G is a function of thebalance between the enzymes systems in the bud, the rate ofsynthesis of the enzymes associated with the development offloral bud parts (floral enzymes) being inversely related tothe level of gibberellins (G) in the bud. The parameter k_G isa function of the nutritional state of the bud; a is a rateparameter, and i is a conversion factor. Node production rateis asymptotic with increase in S, but switches from high tolow values as G varies about a critical concentration. The implicationsof the model, which appears to describe most observations, arediscussed, and some suggestions for testing it are put forward.     

Dieback of rural eucalypts: Response of foliar dietary quality and herbivory to defoliation

Rural dieback of Eucalyptus blakelyi trees growing on pastoral properties near Canberra is associated with chronic defoliation by insects. In order to test the effect of defoliation on subsequent herbivory, I artificially defoliated three healthy trees by clipping their terminal branchlets. The foliage that regrew on the clipped trees was nutritionally superior to the foliage it replaced, and was much more heavily damaged by grazing insects. There was a transient increase in the tannin content of the regrowth foliage, but this was apparently ineffective in defending it from subsequent herbivory. Compared with the foliage on nearby E. blakelyi trees that also produced major flushes of leaf growth during the same period, the regrowth on the clipped trees had enhanced dietary qualities and suffered more insect damage. Leaf age contributed to many of the differences in dietary quality, but when adjustments were made for the effects of leaf age the same trends remained. Five of the nearby trees were suffering from the chronic insect grazing associated with rural dieback, and the other five appeared healthy. The dietary quality of regrowth foliage on the clipped trees was qualitatively more similar to that of foliage on the dieback trees. Thus the chronic herbivory associated with rural dieback may be partly self-perpetuating, given this positive feedback between defoliation and dietary quality, and an apparent absence of other effective controls on insect populations.     

Dieback of rural eucalypts: Does insect herbivory relate to dietary quality of tree foliage?

Foliar dietary quality, and the damage that insects caused to the foliage of dieback-affected and healthy Eucalyptus blakelyi trees, were monitored for 3 years, on pastoral properties in the Australian Capital Territory. Compared with healthy trees, the foliage of dieback trees was more heavily grazed by insects, and its dietary quality was generally superior. Some of the differences in dietary quality were related to the average age of the foliage of healthy and dieback trees. But when statistical models were used to equalize the effects of differences in leaf ages, leaves on dieback trees nonetheless tended to have lower specific weights, and were sometimes rounder and contained more nitrogen. Regression analyses of herbivory against each of the dietary quality variables showed that the only significant relationship that was consistent for both of the years monitored was for foliar nitrogen and herbivory for both dieback and healthy trees. In the first year, regressions between herbivory and specific leaf weight, shape, or sugar content were also significant, but only amongst the dieback trees. This may indicate that these relationships were a response to, rather than a primary cause of, the repeated high defoliation of the dieback trees. Multiple regression equations incorporating annual means of several quality variables explained a high proportion of the variance in annual herbivory, but were grossly different between years.