The Nature and Location of Variable Hydraulic Resistance in Helianthus annuus L. (Sunflower)

Koide, R. 1985. The nature and location of variable hydraulicresistance in Helianthus annuus L. (sunflower).—J. exp.Bot. 36: 1430–1440. Hydraulic resistances for whole sunflower plants (Helianthusannuus L.) and sunflower leaves, stems, petioles and roots weremeasured. Whole plant hydraulic resistance was shown to declinewith an increase in transpiration. Leaf hydraulic resistancewas shown, with one technique employing transpiring leaves,to vary with transpiration and with another technique, employingpressure-induced flow in leaves, to be constant over a widerange of transpiration. Stem and petiole hydraulic resistanceswere constant over a wide range of exudation. Pressure-inducedflow through root systems was shown to be an inappropriate methodfor characterizing their hydraulic properties because flow mayoccur through unnatural paths. The technique employing measuredtranspiration rates and water potentials of non-growing leavesand soil is suggested to be better. The evidence presented inthis study suggests that the hydraulic resistance of the transpirationstream does vary and that the site of variability is the root Key words: Hydraulic resistance, sunflower, pressure-induced water flow     

Modelling of the Hydraulic Architecture of Root Systems: An Integrated Approach to Water Absorption--Distribution of Axial and Radial Conductances in Maize

The ‘Hydraulic Tree Model’ of the root system simulateswater uptake through root systems by coupling a root architecturemodel with laws for water flow into and along roots (Doussan,Pagès and Vercambre,Annals of Botany81: 213–223,1998). A detailed picture of water absorption in all roots comprisingthe root system is thus provided. Moreover, the influence ofdifferent distributions of radial and axial hydraulic conductancesin the root system on the patterns of water uptake can be analysed.Use of the model with Varney and Canny's data (1993) for flowalong maize roots demonstrated that a constant conductance inthe root system cannot reproduce the observed water flux profiles.Taking into account the existing data on hydraulic conductancesin maize roots, we fitted the distribution of conductances inthe root system to the observed flux data. The result is that,during root tissue maturation, the radial conductivity decreasesby one order of magnitude while the axial conductance increasesby about three orders of magnitude. Both types of conductanceexhibit abrupt changes in their evolution. Due to the conductancedistribution in the root system, appreciable water potentialgradients may develop in the roots, in both the branch rootsand main axes. An important point is that the conductance distributionin the branch roots described by the model should be relatedto the age of the tissue (and not the distance from the branchroot tip) and is therefore closely related to the developmentprocess. Thus for branch roots, which represent about 90% ofthe calculated total water uptake in 43-d-old maize, water absorptionwill depend on the opening of the metaxylem in the axes, andon the time dependent variation of the conductances in the branchroots.Copyright 1998 Annals of Botany Company Water; absorption; root system; architecture; model; hydraulic conductance;Zea maysL.     

Effect of genotype and graft type on the hydraulic characteristics and water relations of grafted melon

Abstract

From the measurements of the profiles of hydraulic conductance and water potential from soil through to the leaf system in fully established melon plants, the limits to water flow set by coupling of hydraulic conductance (k) with water relation parameters was evaluated in the laboratory using high pressure flow device (HPFM) and evaporative flux method (EF). The rootstock Arava was grafted onto self, and onto two genotypes (AR57 and AR82) using side and V graft types, and there was an ungrafted control. Hydraulic transport efficiency was estimated from measurements of evaporative flux (transpiration rate) and leaf water potential (ψL) measured between pre-dawn and sunset during the growth cycle. Measured parameters to characterize the hydraulic efficiency (architecture) of the vascular system of melon were normalized to areas of leaves and stem cross section; this enabled the examination of their physiological and ecological functions. The effects of rootstock genotype were more marked on graft union and scion water relations. Differences in the magnitudes of water relation parameters of hydraulic conductance, water potential (lwp) and evaporative water loss (EF) were detected. AR/RS82 side grafted exhibited high EF and Kh despite its lower leaf water potential compared to AR/RS57 V grafted. Self grafting (Arava/Arava grafts) in melon seems to improve water relations and xylem water transport efficiency. Parameters describing the hydraulic efficiency (architecture) of vascular system of melon plants were described in relation to plant attributes. The expression of hydraulic conductance of the root and shoot system relative to plant attributes did not eliminate differences in the magnitudes of conductance elements in tomato and melon. Differences obtained among the different melon grafts in whole plant leaf and stem area specific hydraulic conductance (Kl) indicate the carbon efficiency and hence the cost of resource allocation to areas of root surface and leaves. The role of plant water relations in root-shoot communications and whole plant regulation of water flux are inferred from this study.     

A method for measuring xylem hydraulic conductance and embolism in entire root and shoot systems

Current methods for determining the influence of xylem cavitationon hydraulic conductance are limited to unbranched stem or rootsegments with hydraulic conductances above c. 2 mmol s^–1MPa^–1. Lower conductances and/or highly branched systemsare encountered in seedlings, arid-land shrubs, herbs, and distalportions of shoot and root systems of trees. In order to quantifythe hydraulic impact of cavitation in such systems, existingtechniques have been modified. Branched shoot or root systemswere prepared for measurement by removal of leaves, or roottips, respectively. The shoot or root system was enclosed ina vacuum chamber with the proximal end protruding and suppliedwith perfusing solution. Flow through the xylem was inducedby chamber vacuum. Hydraulic conductance was determined fromthe slope of the flow rate versus pressure relationship. Xylemembolism was quantified from the increase in hydraulic conductancefollowing high pressure (100 kPa) perfusion of solution throughthe plant. Examples are provided of the application of the methodto cavitation studies in the cold desert shrub Artemisia tridentata. Key words: Hydraulic conductance, xylem cavitation, embolism, whole root/shoot system     

Hydraulic resistance components of mature apple trees on rootstocks of different vigours

Dwarfing of fruit trees is often achieved through the use of dwarfing rootstocks. Dwarf trees are characterized by sustained reductions in vegetative growth during the lifetime of the tree. The dwarfing mechanism is not well understood, but it has been hypothesized that hydraulic properties of the rootstock and the graft union are involved. It is hypothesized here that leaf- or stem-specific resistance of at least one hydraulic component of the water transport system would be negatively correlated with rootstock 'vigour', and this could be useful for selection of rootstocks. Hydraulic resistance (R) of fully grown apple trees on a variety of rootstocks of different 'vigours' was measured. Most measurements were with the evaporative flux (EF) method, where water uptake measured with sap flow sensors was related to the pressure gradient from soil (taken as pre-dawn leaf) and midday root (taken as covered root-sucker), stem (from covered leaf), and exposed and shaded leaf water potentials (Psi(l)). R of trees on dwarfing M9 rootstock was compared with that of more vigorous MM106 and MM111 rootstocks in Israel and Vermont, USA. In Israel, M9 consistently had higher leaf-specific hydraulic resistance (R(l)) in the soil to scion stem pathway, but this difference was only significant for one summer. R was larger in M9 between the root and stem, implicating the graft union as the site of increased resistance. In Vermont, R(l) of 9- and 10-year-old trees on six rootstocks of various vigours was not consistently related to vigour, and stem-specific resistance (R(s)) increased with increasing vigour. High pressure flow meter (HPFM) measurements gave a lower R than the EF method in all but one case, perhaps indicating a significant amount of xylem dysfunction in these trees, and demonstrated the increased resistivity of stem sections that included dwarf graft unions as compared with non-graft stem sections. It is concluded that stem- and leaf-specific R are not consistently positively correlated with dwarfing, although the increased resistivity of the graft union in dwarfing rootstocks may influence the transport of water and other elements across the graft union, and therefore be involved in the dwarfing mechanism.     

Shifts in xylem vessel diameter and embolisms in grafted apple trees of differing rootstock growth potential in response to drought

We investigated responses of plant growth rate, hydraulic resistance, and xylem cavitation in scion-rootstock-combinations of Malus domestica L. cv. Honeycrisp scions grafted onto a high-shoot vigor (HSV) rootstock, (semi-dwarfing Malling111), or onto a low-shoot vigor (LSV) rootstock, (dwarfing Budagovsky 9), in response to substrate moisture limitation. Adjustments in xylem vessel diameter and frequency were related to hydraulic resistance measurements for high- versus low- vigor apple trees. We observed a greater tolerance to water deficit in the high-shoot compared to the low-shoot vigor plants under water deficit as evidenced by increased growth in several plant organs, and greater scion anatomical response to limited water availability with ca. 25% increased vessel frequency and ca. 28% narrower current season xylem ring width. Whereas water limitation resulted in greater graft union hydraulic resistance of high-shoot vigor trees, the opposite was true when water was not limiting. The graft union of the low-shoot vigor rootstock exhibited higher hydraulic resistance under well-watered conditions. Scions of high-shoot vigor rootstocks had fewer embolisms at low plant water status compared to scions of low-shoot vigor rootstocks, presumably as a result of large differences in xylem vessel diameter. Our results demonstrated that anatomical differences were related to shifts in hydraulic conductivity and cavitation events, a direct result of grafting, under limited soil water.     

Influence of Soil Water Matric Potential and Hydraulic Conductivity on the Germination of Rape (Brassica napus L.)

The influence of matric potential and hydraulic conductivityon the water absorption and germination of rape (Brassica napusL.) was investigated by comparative studies in three seed-soilwater systems. In all systems the rate of germination and totalgermination were substantially retarded with decreasing matricpotential. Hydraulic conductivity of the soil system was demonstratedto be a limiting factor in the germination process. The effectsof matric potential and hydraulic conductivity on water absorptionfollowed closely those demonstrated for germination. The influenceof matric potential was shown to be comparable to that of osmoticpotential provided consideration was given to the hydraulicproperties of the soil-seed system.     

Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park,Zhejiang Province,China

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.     

浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.     

Modification of Drought Resistance by Water Stress Conditioning in Acacia and Eucalyptus

Plants of Acacia and Eucalyptus species were grown under differentlevels of shading, nutrition, and irrigation to assess the effectof these factors on plant water use. Water use per unit of leaf(phyllode) area was affected only by the irrigation treatment,control plants that had received water daily using appreciablymore water than plants that had been repeatedly subjected towater stress. Water stress conditioning had little or no effecton plant height, leaf (phyllode) area, or minimum stomatal resistancein any of the species. Detailed study of the water stress conditioningof Eucalyptus robusta showed that controls used 46% more waterthan conditioned plants. Leaf area and plant height were unaffectedby conditioning. Control of transpiration was not due to stomatalfunctioning, both sets of plants operating with the same leafdiffusive resistance under conditions of ready water availability.Hydraulic conductivity of the intact root system was loweredby conditioning and it is suggested that this was due, at leastin part, to the effect that conditioning had on root xylem conductivity.Specific conductivity of stem sections was lowered by waterstress conditioning. Water stress avoidance was also associatedwith a more pronounced tendency for stomata to close prior towilting and with a higher level of leaf resistance which couldbe maintained at a low leaf water potential. Conditioned plantsexhibited drought tolerance in their ability to control lossof water from the leaf at lower leaf water potentials than thecontrols.     

叶片结构的水力学特性对植物生理功能影响的研究进展

叶片是植物进行光合、呼吸、蒸腾作用的主要器官, 早期的研究主要集中于水分在叶片中的运输路径, 而对叶脉结构及其生态学意义研究甚少。近年来关于叶片叶脉结构、气孔结构的功能及叶片水力学特性的意义研究已经成为植物生理生态的研究热点。该文综述了叶脉的结构性状的指标(叶脉密度、直径、间距等), 叶片水力学结构特性对植物生长、水分运输、气体交换、光合作用等生理功能的影响, 及其与植物对干旱适应性之间的关系。叶脉结构是决定叶片生理功能的基础, 因此在未来的工作中应分析比较不同种类植物叶脉结构形态与导水、光合、呼吸、同化作用之间的关系, 建立植物茎干-枝-叶系统水力传导的机理性模型, 用以探索不同植物功能结构和高效用水生理生态学机制, 据此评估不同种类植物在未来气候情景下的地位。     

Wound-Induced Hydraulic Signals: Survey of Occurrence in a Range of Species

Localized wounding induces rapid systemic responses in manyplants. These responses include change in surface electricalpotential in various species and induction of proteinase inhibitorgenes in tomato and potato. The nature of the signalling systemwhich co-ordinates such responses is not known, but recently,a role for wound-induced hydraulic signals was proposed. Suchsignals have been reported only from seedlings of wheat andtomato. If they are of any general importance, they must alsooccur in other species. Here, displacement transducers are usedto test for the occurrence of wound-induced hydraulic signalsin a wide range of species. A method is also presented for assessingthe transmission rate in vivo, of the slower component of thewound-induced hydraulic signal (a xylem-borne mass flow of waterand solutes released at the wound site). This component couldbe important in the systemic distribution of elicitors fromsites of localized wounding It is proposed that hydraulic signals could form part of a widespreadmechanism for co-ordination of the plant response to localizedstress Key words: Hydraulic signals, systemic responses, wound-induced effects, ‘PIIF’ transport     

Xylem Hydraulic Characteristics, Water Relations and Wood Anatomy of the Resurrection PlantMyrothamnus flabellifoliusWelw.

Myrothamnus flabellifoliusWelw. is a desiccation-tolerant (‘resurrection’)plant with a woody stem. Xylem vessels are narrow (14 µmmean diameter) and perforation plates are reticulate. This leadsto specific and leaf specific hydraulic conductivities thatare amongst the lowest recorded for angiosperms (k_s0.87 kg m^-1MPa^-1s^-1;k_l3.28x10^-5kg m^-1MPa^-1s^-1, stem diameter 3 mm). Hydraulic conductivitiesdecrease with increasing pressure gradient. Transpiration ratesin well watered plants were moderate to low, generating xylemwater potentials of -1 to -2 MPa. Acoustic emissions indicatedextensive cavitation events that were initiated at xylem waterpotentials of -2 to -3 MPa. The desiccation-tolerant natureof the tissue permits this species to survive this interruptionof the water supply. On rewatering the roots pressures thatwere developed were low (2.4 kPa). However capillary forceswere demonstrated to be adequate to account for the refillingof xylem vessels and re-establishment of hydraulic continuityeven when water was under a tension of -8 kPa. During dehydrationand rehydration cycles stems showed considerable shrinking andswelling. Unusual knob-like structures of unknown chemical compositionwere observed on the outer surface of xylem vessels. These maybe related to the ability of the stem to withstand the mechanicalstresses associated with this shrinkage and swelling.Copyright1998 Annals of Botany Company cavitation, desiccation, hydraulic conductivity, refilling, resurrection plant, root pressure, xylem anatomy,Myrothamnus flabellifolius     

A comparison of plant hydraulic conductances in wheat and lupins

Previous studies have shown similar water use for lupins (Lupinusangustifolius L.) and wheat (Triticum aestivum L.), despitea considerably smaller root system in lupins. A field studyand an experiment under controlled conditions using pressure-fluxrelationships were conducted to examine whether higher hydraulicconductances were responsible for the greater water uptake perunit root length in lupins. In the field experiment, the fluxof water and differences in water potential through the soil-plantsystem were measured for both species and used to calculatethe hydraulic conductance through the plant and through theroot and shoot. The hydraulic conductance for the whole plantwas 3–5 times greater in lupins than in wheat. This relativedifference between the species was similar when plant hydraulicconductance was expressed per unit of root length. This occurreddespite the difference in midday water potential between soiland leaves, being consistently greater in wheat (–1.0MPa) than in lupins (–0.7 MPa). When the total plant conductancewas separated into its components, the combined soil and rootconductance and the shoot conductance were 2 and 6 times greater,respectively, in lupins than in wheat. In the experiment undercontrolled conditions, hydraulic conductance for the entireroot system was determined using a pressure chamber. The specificroot hydraulic conductances were 4 times greater in lupins thanin wheat. The results from both field and controlled conditionsexperiments suggest that the greater water uptake per unit rootlength in lupins compared to wheat results from appreciablylarger root and shoot hydraulic conductances. Key words: Lupins, wheat, hydraulic conductances, water, uptake, pressure-flux     

Water Pathways in Leaves of Hedera helix L. and Tradescantia virginiana L.

Hydraulic conductances of leaf tissues of Hedera helix and Tradescantiavirginiana leaves were measured. It was found that water couldflow most easily through the veins, but that the cell wallsof at least the ventral epidermis were more efficient at resupplyingwater lost from the epidermal tissue than was the mesophyllat rehydrating itself. Vein and bundle-sheath extensions, whichare characteristic of mesomorphic leaves (e.g. T. virginiana),seem to be important in maintaining a close hydraulic connectionbetween the epidermis and the vascular tissue. In leaves notcontaining vein and bundle-sheath extensions, typically xeromorphicleaves (e.g. H. helix), there is not such a close connectionbetween the epidermis and vascular tissue. This was shown inexperiments involving the sudden application of a reduced pressurepotential to either the epidermis or the other tissues of leaves,and the measurement of transient stomatal opening.     

Development and Structure of Unusual Xylem Elements During Graft Union Formation in Picea sitchensis L.

Electron microscope studies of Sitka spruce have been undertakento observe the sequence of cellular development following grafting.Observations at the graft interface reveal a recovery and regenerationsequence associated with union formation. Xylem elements differentiateddirectly from the vascular cambia of the rootstock and scionare different from elements arising from parenchymatous callusderived from ray parenchyma, which may be produced in an attemptto establish a connecting water conducting system as rapidlyas possible. Failure to form such a system may be a primarycause of graft failure. Grafting, Picea sitchensis, xylen elements     

Seasonal Water Acquisition and Redistribution in the Australian Woody Phreatophyte, Banksia prionotes

Sap flows in the xylem of plant roots in response to gradientsin water potential, either between soil and atmosphere (transpiration)or soil layers of different moisture content (termed hydraulicredistribution). The latter has the potential to influence waterbudgets and species interactions, but we lack information forall but a few plant communities. We combined heat pulse measurementsof sap flow with dye and isotope tracing techniques to gaugethe movement of xylem sap within, and exudation from, rootsof Banksia prionotes (Lindley). We demonstrated ‘ hydrauliclift’ during the dry season and provide some evidencethat extremely dry soils limit hydraulic lift. In addition wereport difficulties posed by spiralled xylem tissue in rootsfor the application of heat pulse techniques. Copyright 2000Annals of Botany Company Banksia prionotes, sap flow, hydraulic lift, heat ratio method, deuterium, stable isotopes, root architecture.     

Hydraulic lift among native plant species in the Mojave Desert

Hydraulic lift was investigated among native plants in the Mojave Desert using in situ thermocouple psychrometers. Night lighting and day shading experiments were used to verify the phenomenon. Hydraulic lift was detected for all species examined: five shrub species with different rooting depths and leaf phenologies and one perennial grass species. This study was the first to document hydraulic lift for a CAM species, Yucca schidigera. The pattern of diel flux in soil water potential for the CAM species was temporally opposite to that of C₃ species: for the CAM plant, soil water potential increased in shallow soils during the day when the plant was not transpiring and decreased at night when transpiration began. Because CAM plants transport water to shallow soils during the day when surrounding C₃ and C₄ plants transpire, CAM species that hydraulically lift water may influence water relations of surrounding species to a greater extent than hydraulically lifting C₃ or C₄ species. A strong, negative relationship between the percent sand in the study site soils at the 0.35 m soil depth and the frequency that hydraulic lift was observed at that depth suggests that the occurrence of hydraulic lift is negatively influenced by coarse-textured soils, perhaps due to less root–soil contact in sandy soils relative to finer-textured soils. Differences in soil texture among study sites may explain, in part, differences in the frequency that hydraulic lift was detected among these species. Further investigations are needed to elucidate species versus soil texture effects on hydraulic lift. This revised version was published online in June 2006 with corrections to the Cover Date.