The compensation heat-pulse method for measuring sap flow is tested here in olive trees (Olea europaea L.). We describe a rigorous three-way examination of the robustness of the technique for this species, and examine the potential of the technique for an automatic control of the irrigation system. Two tests were carried out using heat-pulse gear inserted into the stem of 12-year-old 'Manzanilla' olive trees. One test used forced-flow through a stem section, and the other involved measured water uptake by an excised tree. The measured sap flow in these two tests was in agreement with calculations from heat-pulse velocities when using a standard 'wound correction' to account for the presence of the probes and the disruption to the sap flow. Thus, this technique for monitoring transpiration can, we feel, be used with confidence in olives. The third experiment was carried out in the field , where we analysed sap flow data from two 29year-old olive trees-one tree was under regular drip irrigation and the other was from dry-farming conditions. We use measurements of sap flow in the trunk to examine the hydraulic functioning of the tree, and to explore sorne diagnostics of water stress. Our heat-pulse measurements in the irrigated olive tree exhibited a profile of sap flow that was weighted towards the outer xylem of the tree trunk while the water-stressed trees in the field showed a profile of sap flow weighted towards the centre of the trunk. The loss of hydraulic functioning in the outermost section of the vascular system, as a result of water stress, we consider to be due both to stomatal control and to embolisms in the xylem vessels. The fourth experiment was also carried out in the field, in which sap flow measurements were made at three locations in the trunk as well as in two roots of another 29-year-old olive tree. The soil explored by each root, on opposite sides of the trunk, was differentially wetted by separate irrigation of each side. Our data showed that the surface roots were able to absorb water immediately after wetting, despite a reasonably prolonged period of moderate drought. Root activity quickly shifted to the regions where the soil had been wetted. A root in dry soil exhibited no flow at night, whereas sap flows of about 0.02 l h À1 were measured around midnight in the root drawing water from the wetter soil. Our observations suggest that the hydraulic behaviour of the trunk and surface roots might be used as a diagnostic of the onset, or severity, of water stress. Here there is not the imperative to replicate, for the prime goal is not transpiration estimation. Rather interpretation of the diurnal dynamics is used to infer the onset, or severity of water stress. The compensation heat-pulse seems a suitable technique for automatically controlling the irrigation system of olives, and probably other trees, based either on the estimation of the short-time dynamics of transpiration, or on changes in the hydraulic behaviour of the trees.
It is widely believed that partial root drying (PRD) reduces water losses by transpiration without affecting yield. However, experimental work carried out to date does not always support this hypothesis. In many cases a PRD treatment has been compared to a full irrigated treatment, so doubt remains on whether the observed benefits correspond to the switching of irrigation or just to PRD being a deficit irrigation treatment. In addition, not always a PRD treatment has been found advantageous as compared to a companion regulated deficit irrigation (RDI) treatment. In this work we have compared the response of mature 'Manzanilla' olive trees to a PRD and an RDI treatment in which about 50% of the crop evapotranspiration (ET c ) was supplied daily by localised irrigation. We alternated irrigation in the PRD treatment every 2 weeks in 2003 and every 3 weeks in 2004. Measurements of stem water potential (Y stem ), stomatal conductance (g s ) and net CO 2 assimilation rate (A) were made in trees of both treatments, as well as in trees irrigated to 100% of ET c (Control trees) and in Rain-fed trees. Sap flow was also measured in different conductive organs of trees under both PRD and RDI treatments, to evaluate the influence of alternating irrigation on root water uptake and tree water consumption. We found small and random differences in Y stem , g s and A, which gave no evidence of PRD causing a positive effect on the olive tree performance, as compared to RDI. Stomatal conductance decreased in PRD trees as compared to Control trees, but a similar decrease in g s was also recorded in the RDI trees. Sap flow measurements, which reflected water use throughout the irrigation period, also showed no evidence of g s being more reduced in PRD than in RDI trees. Daily water consumption was also similar in the trees of the deficit irrigation treatments, for most days, throughout the irrigation period. Alternating irrigation in PRD trees did not cause a change in either water taken up by main roots at each side of the trees, or in the sap flow of both trunk locations and main branches of each side. Results from this work, and from previous work conducted in this orchard, suggest that transpiration is restricted in trees under deficit irrigation, in which roots are left in drying soil when water is applied by localised irrigation, and that there is no need to alternate irrigation for achieving this effect.
We quantified parameters for a model of leaf-level photosynthesis for olive, and tested the model against an independent dataset. Specific temperature-dependence parameters of the model for olive leaves were measured, as well as the relationship of the model parameters with area-based leaf nitrogen (N) content. The effect of soil water deficit on leaf photosynthesis was examined by applying two irrigation treatments to 29-year-old trees growing in a plantation: drip irrigation sufficient to meet the crop water requirements (I) and dry-farming (D). In both treatments, leaves had a higher photosynthetic capacity in April than in August. In August, photosynthetic capacity was lower in D trees than in I trees. Leaf photosynthetic capacity was linearly and positively related to leaf N content on an area basis (N(a)) and to leaf mass per unit area (LMA), and the regression slope varied with irrigation treatment. The seasonal reduction in N(a) was used in the model to predict photosynthesis under drought conditions. Olive leaves showed a clear limitation of photosynthesis by triose phosphate utilization (TPU) even at 40 degrees C, and the data suggest that olive invests fewer resources in TPU than other species. The seasonal decrease in photosynthetic capacity moderated the stomatal limitation to carbon dioxide (CO(2)) fixation as soil water deficit increased. Further, it enabled leaves to operate close to the transition point between photosynthetic limitation due to RuBP carboxylation capacity and that due to RuBP regeneration capacity, and resulted in a near constant value of internal CO(2) concentration from April to August. Under well watered conditions, N-use efficiency of the olive leaves was enhanced at the expense of reduced water-use efficiency.
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