Xylem Flow Research Articles

Water Transport Properties of the Grape Pedicel during Fruit Development: Insights into Xylem Anatomy and Function Using Microtomography.

Xylem flow of water into fruits declines during fruit development, and the literature indicates a corresponding increase in hydraulic resistance in the pedicel. However, it is unknown how pedicel hydraulics change developmentally in relation to xylem anatomy and function. In this study on grape (Vitis vinifera), we determined pedicel hydraulic conductivity (kh) from pressure-flow relationships using hydrostatic and osmotic forces and investigated xylem anatomy and function using fluorescent light microscopy and x-ray computed microtomography. Hydrostatic kh (xylem pathway) was consistently 4 orders of magnitude greater than osmotic kh (intracellular pathway), but both declined before veraison by approximately 40% and substantially over fruit development. Hydrostatic kh declined most gradually for low (less than 0.08 MPa) pressures and for water inflow and outflow conditions. Specific kh (per xylem area) decreased in a similar fashion to kh despite substantial increases in xylem area. X-ray computed microtomography images provided direct evidence that losses in pedicel kh were associated with blockages in vessel elements, whereas air embolisms were negligible. However, vessel elements were interconnected and some remained continuous postveraison, suggesting that across the grape pedicel, a xylem pathway of reduced kh remains functional late into berry ripening.

Living cells in wood. 1. Absence, scarcity and histology of axial parenchyma as keys to function

The diversity of expression in axial parenchyma (or lack of it) in woods is reviewed and synthesized with recent work in wood physiology, and questions and hypotheses relative to axial parenchyma anatomy are offered. Cell shape, location, abundance, size, wall characteristics and contents are all characteristics for the assessment of the physiological functions of axial parenchyma, a tissue that has been neglected in the consideration of how wood histology has evolved. Axial parenchyma occurrence should be considered with respect to mechanisms for the prevention and reversal of embolisms in tracheary elements. This mechanism complements cohesion–tension-based water movement and root pressure as a way of maintaining flow in xylem. Septate fibres can substitute for axial parenchyma (‘axial parenchyma absent’) and account for water movement in xylem and for the supply of carbohydrate abundance underlying massive and sudden events of foliation, flowering and fruiting, as can fibre dimorphism and the co-occurrence of septate fibres and axial parenchyma. Rayless woods may or may not contain axial parenchyma and are informative when analysing parenchyma function. Interconnections between ray and axial parenchyma are common, and so axial and radial parenchyma must be considered as complementary parts of a network, with distinctive but interactive functions. Upright ray cells and more numerous rays per millimetre enhance interconnection and are more often found in woods that contain tracheids. Vesselless woods in both gymnosperms and angiosperms have axial parenchyma, the distribution of which suggests a function in osmotic water shifting. Water and photosynthate storage in axial parenchyma may be associated with seasonal changes and with succulent or subsucculent modes of construction. Apotracheal axial parenchyma distribution often demonstrates storage functions that can be read independently of osmotic water shifting capabilities. Axial parenchyma may serve to both enhance mechanical strength or, when parenchyma is thin-walled, as a tissue that adapts to volume change with a change in water content. Other functions of axial parenchyma (contributing resistance to pathogens; a site for the recovery of physical damage) are considered. The diagnostic features of axial parenchyma and septate fibres are reviewed in order to clarify distinctions and to aid in cell type identification. Systematic listings are given for particular axial parenchyma conditions (e.g. axial parenchyma ‘absent’ with septate fibres substituting). A knowledge of the axial parenchyma information presented here is desirable for a full understanding of xylem function.

The decline in xylem flow to mango fruit at the end of its development is related to the appearance of embolism in the fruit pedicel.

The decline in xylem flow during the late growth stage in most fruits may be due either to a decrease in the water potential gradient between the stem bearing the fruit and the fruit tissues or to a decrease in the hydraulic conductivity of xylem vessels, or both. In this study, we analysed changes in xylem flows to the mango Mangifera indica L. fruit during its development to identify the sources of variation by measuring changes in the water potential gradient and in the hydraulic properties of the fruit pedicel. The variations in xylem and transpiration flows were estimated at several stages of mango fruit development from the daily changes in the fresh mass of detached and girdled fruits on branches. The water potential gradient was estimated by monitoring the diurnal water potential in the stem and fruit. The hydraulic properties of the fruit pedicel were estimated using a flow meter. The results indicated that xylem flow increased in the early stages of fruit development and decreased in the late stage. Variations in xylem flow were related to the decrease in the hydraulic conductivity of xylem vessels but not to a decrease in the water potential gradient. The hydraulic conductivity of the fruit pedicel decreased during late growth due to embolism caused by a decrease in the fruit water potential. Further studies should establish the impact of the decrease in the hydraulic conductivity of the fruit pedicel on mango growth.

Mathematical modeling of positron emission tomography (PET) data to assess radiofluoride transport in living plants following petiolar administration.

BackgroundIon transport is a fundamental physiological process that can be studied non-invasively in living plants with radiotracer imaging methods. Fluoride is a known phytotoxic pollutant and understanding its transport in plants after leaf absorption is of interest to those in agricultural areas near industrial sources of airborne fluoride. Here we report the novel use of a commercial, high-resolution, animal positron emission tomography (PET) scanner to trace a bolus of [18F]fluoride administered via bisected petioles of Brassica oleracea, an established model species, to simulate whole plant uptake of atmospheric fluoride. This methodology allows for the first time mathematical compartmental modeling of fluoride transport in the living plant. Radiotracer kinetics in the stem were described with a single-parameter free- and trapped-compartment model and mean arrival times at different stem positions were calculated from the free-compartment time-activity curves.ResultsAfter initiation of administration at the bisected leaf stalk, [18F] radioactivity climbed for approximately 10 minutes followed by rapid washout from the stem and equilibration within leaves. Kinetic modeling of transport in the stem yielded a trapping rate of 1.5 +/− 0.3%/min (mean +/− s.d., n = 3), velocity of 2.2 +/− 1.1 cm/min, and trapping fraction of 0.8 +/− 0.5%/cm.ConclusionQuantitative assessment of physiologically meaningful transport parameters of fluoride in living plants is possible using standard positron emission tomography in combination with petiolar radiotracer administration. Movement of free fluoride was observed to be consistent with bulk flow in xylem, namely a rapid and linear change in position with respect to time. Trapping, likely in the apoplast, was observed. Future applications of the methods described here include studies of transport of other ions and molecules of interest in plant physiology.Electronic supplementary materialThe online version of this article (doi:10.1186/s13007-015-0061-y) contains supplementary material, which is available to authorized users.

Sugar demand of ripening grape berries leads to recycling of surplus phloem water via the xylem.

We tested the common assumption that fleshy fruits become dependent on phloem water supply because xylem inflow declines at the onset of ripening. Using two distinct grape genotypes exposed to drought stress, we found that a sink-driven rise in phloem inflow at the beginning of ripening was sufficient to reverse drought-induced berry shrinkage. Rewatering accelerated berry growth and sugar accumulation concurrently with leaf photosynthetic recovery. Interrupting phloem flow through the peduncle prevented the increase in berry growth after rewatering, but interrupting xylem flow did not. Nevertheless, xylem flow in ripening berries, but not berry size, remained responsive to root or shoot pressurization. A mass balance analysis on ripening berries sampled in the field suggested that phloem water inflow may exceed growth and transpiration water demands. Collecting apoplastic sap from ripening berries showed that osmotic pressure increased at distinct rates in berry vacuoles and apoplast. Our results indicate that the decrease in xylem inflow at the onset of ripening may be a consequence of the sink-driven increase in phloem inflow. We propose a conceptual model in which surplus phloem water bypasses the fruit cells and partly evaporates from the berry surface and partly moves apoplastically to the xylem for outflow.

Leaf gas exchanges and water relations affect the daily patterns of fruit growth and vascular flows in Abbé Fétel pear (Pyrus communis L.) trees

Daily patterns of leaf and stem water potentials, leaf gas exchanges (net photosynthesis, stomatal conductance and transpiration), fruit growth, vascular (phloem and xylem) and transpiration in/outflows were determined in two subsequent years at regular time intervals during the season on pear trees of the cv. Abbé Fétel. Leaf gas exchanges showed higher values in the morning, while fruit shrank due to high transpiration water losses which were not balanced by xylem flows. Despite the high rates of transpiration, fruit were fully rehydrated in the afternoon thanks to high xylem flows, while phloem import occurred mostly in the midday hours. During most of the season, fruit growth was sustained above all by the xylem flux which represented upto 85% of fruit total inflows. Later, fruit growth rate decreased while dry matter accumulation increased following a rise in the phloem relative contribution to growth, which reached 50% of fruit total inflows. This is the first report to show how the biophysical mechanism of fruit growth in “Abbé Fétel” pear is based on high water exchanges by xylem and transpiration during most of the season. The simultaneous monitoring of leaf gas exchanges and fruit vascular flows during the day suggests how until about 95 days after full bloom (DAFB), leaves are stronger “sinks” for water in the morning while fruit import water mainly during the afternoon, thanks to the increase in leaf water potential following stomatal closure.

Phloem flow and sugar transport in Ricinus communis L. is inhibited under anoxic conditions of shoot or roots.

Anoxic conditions should hamper the transport of sugar in the phloem, as this is an active process. The canopy is a carbohydrate source and the roots are carbohydrate sinks. By fumigating the shoot with N2 or flooding the rhizosphere, anoxic conditions in the source or sink, respectively, were induced. Volume flow, velocity, conducting area and stationary water of the phloem were assessed by non-invasive magnetic resonance imaging (MRI) flowmetry. Carbohydrates and δ(13) C in leaves, roots and phloem saps were determined. Following flooding, volume flow and conducting area of the phloem declined and sugar concentrations in leaves and in phloem saps slightly increased. Oligosaccharides appeared in phloem saps and after 3 d, carbon transport was reduced to 77%. Additionally, the xylem flow declined and showed finally no daily rhythm. Anoxia of the shoot resulted within minutes in a reduction of volume flow, conductive area and sucrose in the phloem sap decreased. Sugar transport dropped to below 40% by the end of the N2 treatment. However, volume flow and phloem sap sugar tended to recover during the N2 treatment. Both anoxia treatments hampered sugar transport. The flow velocity remained about constant, although phloem sap sugar concentration changed during treatments. Apparently, stored starch was remobilized under anoxia.

Increasing water stress negatively affects pear fruit growth by reducing first its xylem and then its phloem inflow

Drought stress negatively affects many physiological parameters and determines lower yields and fruit size. This paper investigates on the effects of prolonged water restriction on leaf gas exchanges, water relations and fruit growth on a 24-h time-scale in order to understand how different physiological processes interact to each other to face increasing drought stress and affect pear productive performances during the season. The diurnal patterns of tree water relations, leaf gas exchanges, fruit growth, fruit vascular and transpiration flows were monitored at about 50, 95 and 145 days after full bloom (DAFB) on pear trees of the cv. Abbé Fétel, subjected to two irrigation regimes, corresponding to a water restitution of 100% and 25% of the estimated Etc, respectively. Drought stress progressively increased during the season due to lower soil tensions and higher daily vapour pressure deficits (VPDs). Stem water potential was the first parameter to be negatively affected by stress and determined the simultaneous reduction of fruit xylem flow, which at 95 DAFB was reflected by a decrease in fruit daily growth. Leaf photosynthesis was reduced only from 95 DAFB on, but was not immediately reflected by a decrease in fruit phloem flow, which instead was reduced only at 145 DAFB. This work shows how water stress negatively affects pear fruit growth by reducing first its xylem and then its phloem inflow. This determines a progressive increase in the phloem relative contribution to growth, which lead to the typical higher dry matter percentages of stressed fruit.

Uptake and distribution of chlordecone in radish: Different contamination routes in edible roots

Chlordecone (CLD) was an organochlorine insecticide mainly used to struggle against banana weevils in the French West Indies. Forbidden since 1993, it has been a long-term contaminant of soils and aquatic environments. Crops growing in contaminated soils lead to human exposure by food consumption. We used radiolabeled [14C]-CLD to investigate the contamination ways into radish, a model of edible roots. Radish plants were able to accumulate CLD in both roots (RCF35d 647) and tubers (edible parts, CF35d 6.3). CLD was also translocated to leaves (CF35d 1.7). The contamination of tuber was mainly due to peridermic adsorption or CLD systemic translocation to the pith. TSCF was 3.44×10−3. CLD diffused across periderm to internal tissues. We calculated a mean flux of diffusion J through periderm about 5.71×10−14gcm−2s−1.We highlighted different contamination routes of the tuber, (i) adsorption on periderm followed by diffusion of CLD towards underlying tissues, cortex, xylem, and pith (ii) adsorption by roots and translocation by the transpiration stream followed by diffusion from xylem vessels towards inner tissues, pith, and peripheral tissues, cortex and periderm. Concerning chemical risk assessment for other tubers, contamination would depend on various parameters, the thickness of periderm and CLD periderm permeance, the origin of secondary tissues – from cortex and/or pith – , the importance of xylem flow in tuber, and the lipid amount within tuber.

Significance of xylem water conductance for the compatibility of maté phenotypes (Ilex paraguarensis) for grafting

In this study the compatibility for grafting of root stocks and buds of three recognized leaf phenotypes of Ilex paraguarensis St. (mate) Hil. from south Brazil was investigated. Special regard was given to the anatomical structure and the sap flow in the secondary xylem of the grafted plants. Significant differences of the wood structure expressed in terms of vessel portion, vessel member length, and intervessel perforations were found in low compatible phenotypes, while highly compatible phenotypes had a very similar wood structure. In grafts of highly compatible phenotypes six months after grafting the water conducting system of root stocks and buds was strongly linked to each other, while in grafts of low compatible phenotypes the formation of connecting vessel elements was rare. In the first month after grafting the xylem sap flow in the buds of all grafts was significantly higher compared to their root stocks, while in the second month in grafts of highly compatible phenotypes, the water transport in the root stocks and in the buds was balanced. In contrast, in grafts of low compatible phenotypes even in the second month the water transport in the root stocks was lower compared to the buds causing the dieback of a high portion of these plants.

Tree shoot bending generates hydraulic pressure pulses: a new long-distance signal?

When tree stems are mechanically stimulated, a rapid long-distance signal is induced that slows down primary growth. An investigation was carried out to determine whether the signal might be borne by a mechanically induced pressure pulse in the xylem. Coupling xylem flow meters and pressure sensors with a mechanical testing device, the hydraulic effects of mechanical deformation of tree stem and branches were measured. Organs of several tree species were studied, including gymnosperms and angiosperms with different wood densities and anatomies. Bending had a negligible effect on xylem conductivity, even when deformations were sustained or were larger than would be encountered in nature. It was found that bending caused transient variation in the hydraulic pressure within the xylem of branch segments. This local transient increase in pressure in the xylem was rapidly propagated along the vascular system in planta to the upper and lower regions of the stem. It was shown that this hydraulic pulse originates from the apoplast. Water that was mobilized in the hydraulic pulses came from the saturated porous material of the conduits and their walls, suggesting that the poroelastic behaviour of xylem might be a key factor. Although likely to be a generic mechanical response, quantitative differences in the hydraulic pulse were found in different species, possibly related to differences in xylem anatomy. Importantly the hydraulic pulse was proportional to the strained volume, similar to known thigmomorphogenetic responses. It is hypothesized that the hydraulic pulse may be the signal that rapidly transmits mechanobiological information to leaves, roots, and apices.

Tree root systems competing for soil moisture in a 3D soil–plant model

Competition for water among multiple tree rooting systems is investigated using a soil–plant model that accounts for soil moisture dynamics and root water uptake (RWU), whole plant transpiration, and leaf-level photosynthesis. The model is based on a numerical solution to the 3D Richards equation modified to account for a 3D RWU, trunk xylem, and stomatal conductances. The stomatal conductance is determined by combining a conventional biochemical demand formulation for photosynthesis with an optimization hypothesis that selects stomatal aperture so as to maximize carbon gain for a given water loss. Model results compare well with measurements of soil moisture throughout the rooting zone, of total sap flow in the trunk xylem, as well as of leaf water potential collected in a Loblolly pine forest. The model is then used to diagnose plant responses to water stress in the presence of competing rooting systems. Unsurprisingly, the overlap between rooting zones is shown to enhance soil drying. However, the 3D spatial model yielded transpiration-bulk root-zone soil moisture relations that do not deviate appreciably from their proto-typical form commonly assumed in lumped eco-hydrological models. The increased overlap among rooting systems primarily alters the timing at which the point of incipient soil moisture stress is reached by the entire soil–plant system.

Vigour reduction in girdled peach trees is related to lower midday stem water potentials

Stem or trunk girdling is a technique used in physiological studies and in horticultural practice for interrupting carbon flow through the phloem to other parts of the plant without influencing water flow in the xylem. Trunk girdling in peaches is practiced primarily to stimulate fruit growth but it also tends to decrease shoot vigour for a period of time after girdling. Water flow through the trunk or branches of peach trees is thought to be primarily dependent on the most recently formed ring of xylem and vegetative growth is closely related to stem water potential and stem hydraulic conductance. The aim of the present work was to determine whether vigour reduction due to girdling was correlated with a reduction in midday stem water potential during the period of time between girdling and the subsequent healing of stem tissue. 'Springcrest' peach trees were girdled on two different dates. Fruit yield and size, water sprout growth, proleptic shoot growth and stem water potential were measured. Early and late girdled trees yielded larger fruits and fewer and shorter water sprouts in comparison with control trees. Midday stem water potential declined significantly after girdling and gradually recovered until the time of fruit harvest. These results suggest that the vigour reduction of girdled trees is related to a decrease of midday stem water potential caused by girdling. Early tree girdling increased the reduction in midday stem water potential and shoot growth compared with the later girdling treatment. These results point out that even though girdling only removes bark and phloem tissue it can apparently affect water flow in xylem.

Dynamic State of Water in Excised Ligustrum Lucidum Branches Observed by Dedicated Micro-Magnetic Resonance Imaging

The dynamic state of water was investigated for tree branches by a combined k-space and q-space imaging method using a dedicated magnetic resonance imaging (MRI) device equipped with a 1.0-T permanent magnet. Changes in the 1H-NMR image due to the application of pulsed-field gradients (PFG) of stepping magnitude were measured using a phantom consisting of flow water and stationary water; the relationship between the flow velocities and the diffusion coefficients of water was studied. The method was applied to excised Ligustrum lucidum branches under conditions with and without weak light (100-140 μmol m-2 s-1). The distributions and diffusion coefficients of cell-associated water for individual tissues in the branches were determined in relation to morphology. Large amounts of water existed but diffusion coefficients were not high in the cambium. Though the water amounts were moderate, the highest diffusion coefficient exceeded that for pure water in the secondary xylem. The conduit tubes were smaller than the imaging pixel, unlike the phantom, so the determined values might be perturbed by the conditions of the ambient cell-associated water. However, positional responses in the xylem flow to light were detected, and a flow velocity up to 0.71 mm s-1 by transpiration was recorded. The movement of water in the phloem was not distinguished from large amounts of water in adjacent tissues, probably due to the very small cells and slow rates of flow in the phloem.

A closed‐form solution for steady‐state coupled phloem/xylem flow using the Lambert‐W function

A closed-form solution for steady-state coupled phloem/xylem flow is presented. This incorporates the basic Münch flow model of phloem transport, the cohesion model of xylem flow, and local variation in the xylem water potential and lateral water flow along the transport pathway. Use of the Lambert-W function allows this solution to be obtained under much more general and realistic conditions than has previously been possible. Variation in phloem resistance (i.e. viscosity) with solute concentration, and deviations from the Van't Hoff expression for osmotic potential are included. It is shown that the model predictions match those of the equilibrium solution of a numerical time-dependent model based upon the same mechanistic assumptions. The effect of xylem flow upon phloem flow can readily be calculated, which has not been possible in any previous analytical model. It is also shown how this new analytical solution can handle multiple sources and sinks within a complex architecture, and can describe competition between sinks. The model provides new insights into Münch flow by explicitly including interactions with xylem flow and water potential in the closed-form solution, and is expected to be useful as a component part of larger numerical models of entire plants.

Hydraulic resistance of developing Actinidia fruit

Xylem flows into most fruits decline as the fruit develop, with important effects on mineral and carbohydrate accumulation. It has been hypothesized that an increase in xylem hydraulic resistance (RT) contributes to this process. This study examined changes in RT that occur during development of the berry of kiwifruit (Actinidia deliciosa), identified the region within the fruit where changes were occurring, and tested whether a decrease in irradiance during fruit development caused an increase in RT, potentially contributing to decreased mineral accumulation in shaded fruit. RT was measured using pressure chamber and flow meter methods, the two methods were compared, and the flow meter was also used to partition RT between the pedicel, receptacle and proximal and distal portions of the berry. Dye was used as a tracer for xylem function. Artificial shading was used to test the effect of light on RT, dye entry and mineral accumulation. RT decreased during the early phase of rapid fruit growth, but increased again as the fruit transitioned to a final period of slower growth. The most significant changes in resistance occurred in the receptacle, which initially contributed 20 % to RT, increasing to 90 % later in development. Dye also ceased moving beyond the receptacle from 70 d after anthesis. The two methods for measuring RT agreed in terms of the direction and timing of developmental changes in RT, but pressure chamber measurements were consistently higher than flow meter estimates of RT, prompting questions regarding which method is most appropriate for measuring fruit RT. Shading had no effect on berry growth but increased RT and decreased dye movement and calcium concentration. Increased RT in the receptacle zone coincides with slowing fresh weight growth, reduced transpiration and rapid starch accumulation by the fruit. Developmental changes in RT may be connected to changes in phloem functioning and the maintenance of water potential gradients between the stem and the fruit. The effect of shade on RT extends earlier reports that shading can affect fruit vascular differentiation, xylem flows and mineral accumulation independently of effects on transpiration.

LIMITATIONS IN THE HYDRAULIC PATHWAY: EFFECTS OF XYLEM EMBOLISMS ON SAP VELOCITY AND FLOW

LIMITATIONS IN THE HYDRAULIC PATHWAY: EFFECTS OF XYLEM EMBOLISMS ON SAP VELOCITY AND FLOW

Multiple impacts of the plant growth-promoting rhizobacterium Variovorax paradoxus 5C-2 on nutrient and ABA relations of Pisum sativum.

Resolving the physiological mechanisms by which rhizobacteria enhance plant growth is difficult, since many such bacteria contain multiple plant growth-promoting properties. To understand further how the 1-aminocyclopropane-1-carboxylate (ACC) deaminase (ACCd)-containing rhizobacterium Variovorax paradoxus 5C-2 affects plant growth, the flows and partitioning of mineral nutrients and abscisic acid (ABA) and ABA metabolism were studied in pea (Pisum sativum) plants following rhizosphere bacterial inoculation. Although root architecture was not affected, inoculation increased root and shoot biomass, and stomatal conductance, by 20, 15, and 24%, respectively, and increased N, P, K, Ca, and Mg uptake by 16, 81, 50, 46, and 58%, respectively. P deposition in inoculated plant roots was 4.9 times higher than that in uninoculated controls. Rhizobacterial inoculation increased root to shoot xylem flows and shoot to root phloem flows of K by 1.8- and 2.1-fold, respectively. In control plants, major sinks for K deposition were the roots and upper shoot (43% and 49% of total uptake, respectively), while rhizobacterial inoculation increased K distribution to the lower shoot at the expense of other compartments (xylem, phloem, and upper shoot). Despite being unable to metabolize ABA in vitro, V. paradoxus 5C-2 decreased root ABA concentrations and accumulation by 40–60%. Although inoculation decreased xylem ABA flows, phloem ABA flows increased. Whether bacterial ACCd attenuates root to shoot ABA signalling requires further investigation, since ABA is critical to maintain growth of droughted plants, and ACCd-containing organisms have been advocated as a means of minimizing growth inhibition of plants in drying soil.

Short-period changes in weather conditions affect xylem, but not phloem flows to young kiwifruit (Actinidia deliciosa) berries

Weather conditions are known to affect many physiological parameters like canopy gas exchanges, tree water relations and fruit growth. However, their effects may be different depending on plant species and developmental stage. This study investigates the effects of sudden weather worsening on the daily dynamics of kiwifruit berry growth, at different times during the season. Phloem, xylem and transpiration flows to/from the berry were continuously monitored for several days at about 5, 9, 11 and 15 weeks after full bloom (WAFB), on two consecutive years, while air temperature, relative humidity, solar radiation and rain were simultaneously recorded. From these data vapour pressure deficit (VPD) was also calculated. For each period considered, the daily amounts and the daily patterns of kiwifruit berry relative growth rate (RGR) and vascular flows were compared between two subsequent days having high (HVPD) and low (LVPD) mean VPDs, respectively. During all periods, daily water exchanges were linearly related to VPD although these relationships showed decreasing slopes during the season. At 5 and 9 WAFB daily and hourly rates of xylem and transpiration flows were highly reduced by LVPD conditions, while phloem flow was not affected by such changes during the whole season. Also, LVPD conditions affected the typical daily fluctuations in the berry RGR by reducing its morning shrinkage and slowing down its afternoon growth rates. This study shows how in the early stages of kiwifruit berry growth, weather worsening highly reduces the berry water exchanges by xylem and transpiration, but does not affect its phloem imports in the short-period.

Vascular functioning and the water balance of ripening kiwifruit (Actinidia chinensis) berries.

Indirect evidence suggests that water supply to fleshy fruits during the final stages of development occurs through the phloem, with the xylem providing little water, or acting as a pathway for water loss back to the plant. This inference was tested by examining the water balance and vascular functioning of ripening kiwifruit berries (Actinidia chinensis var. chinensis ‘Hort16A’) exhibiting a pre-harvest ‘shrivel’ disorder in California, and normal development in New Zealand. Dye labelling and mass balance experiments indicated that the xylem and phloem were both functional and contributed approximately equally to the fruit water supply during this stage of development. The modelled fruit water balance was dominated by transpiration, with net water loss under high vapour pressure deficit (Da) conditions in California, but a net gain under cooler New Zealand conditions. Direct measurement of pedicel sap flow under controlled conditions confirmed inward flows in both the phloem and xylem under conditions of both low and high Da. Phloem flows were required for growth, with gradual recovery after a step increase in Da. Xylem flows alone were unable to support growth, but did supply transpiration and were responsive to Da-induced pressure fluctuations. The results suggest that the shrivel disorder was a consequence of a high fruit transpiration rate, and that the perception of complete loss or reversal of inward xylem flows in ripening fruits should be re-examined.