The Development of the Runner Bean Leaf with Special Reference to the Relation between the Sizes of the Lamina and of the Petiolar Xylem: III. The Development of the Leaf under Various Conditions

The development of the first pair of leaves of the Runner Bean(Phaseolus multiflorus Willd.) under various conditions is described. 1. Under damp conditions and also when one leaf of a pair wascovered with a black paper envelope, the rate of developmentwas altered but no significant changes in the relative developmentof the parts of the leaf occurred. 2. Removal of a portion of the lamina at an early stage of developmentresulted, at maturity, in a leaf having relatively less petiolarxylem, although the ratio xylem area/lamina area was slightlyhigher. This treatment also delayed a ‘sudden’ increasein area of the vessels which was seen in normal plants. 3. An attempt was made to estimate the transporting power ofthe petiolar xylem and to compare it with the lamina area. Thedata suggest that the maximum possible area of the lamina atany stage in development may be determined by the transportingpower of the xylem, but that the attainment of this maximumpossible area may be prevented by external factors as is seenin some of the experimental results presented.     

The Development of the Runner-bean Leaf with Special Reference to the Relation between the Sizes of the Lamina and of the Petiolar Xylem: II. The Normal Development of the Bean Leaf

The development of the first pair of leaves of the Runner-bean(Phaseolus multiflorus Willd.) has been investigated from theunfolding of the leaves to the end of lamina expansion. Thegrowth of the petiole is relatively fast and ceases first. Extensiongrowth of the lamina continues longer. In the growth of thexylem, the average area of the vessels formed increases mostmarkedly at an early developmental stage. Vessel number maycontinue to increase after lamina growth has ceased. Both thenumber of vessels and the xylem area per unit area of laminadiminish as the leaf matures.     

The Initiation and Development of Secondary Xylem in Axillary Branches of Populus deltoides

The initiation of secondary xylem in elongating axillary branchesof Populus deltoides Bartr. ex Marsh. is independent of thatin the main stem. Although secondary xylem differentiates acropetallyin the main stem, it does not differentiate from the stem intothe axillary branch. Secondary xylem is usually initiated ininternode 4 (occasionally 3) of the axillary branch, and fromthis site it develops both acropetally in the elongating branchand basipetally toward the main stem. Secondary vessel differentiationalways precedes fibre differentiation. Although secondary xylemdifferentiates in internodes that have ceased elongation, itdifferentiates first in traces of the vascular cylinder servingrapidly expanding and maturing foliage leaves. As younger leaveson the branch expand and mature, secondary xylem differentiatesin their traces eventually producing a complete secondary vascularcylinder. Scale leaves do not initiate secondary xylem independentlyin their traces; they are activated by adjacent traces in thevascular cylinder serving foliage leaves. Once established,the primary-secondary vascular transition zone advances acropetallyin a branch just as it does in the main stem. Populus deltoides Bartr. ex Marsh., cottonwood, axillary branches, secondary xylem, plastochron index, post-dormancy development, xylem.     

Ionic Interactions of Petiole and Lamina During the Life of a Leaf of Castor Bean (Ricinus communis L.) Under Moderately Saline Conditions

Ion (K⁺, Na⁺, Mg²⁺, Ca²⁺ and Cl) flows and partitioning in thepetiole and lamina of leaf 6 of castor bean {Ricinus communisL.) plants growing in the presence of a mean of 71 mol m^–3NaCl were described by an empirical modelling technique. Thiscombined data on changes in ion contents of petiole and lamina,ion: carbon molar ratios of phloem bleeding sap and pressure-inducedxylem exudates of the leaf with previously described informationon the economies of C and N in identical leaf material. Datawere expressed as daily exchanges of ions in xylem and phloem,or depicted as models of ion balance and transport activityof petiole and lamina during four consecutive phases of leaflife. The early import phase was characterized by high intakeof K and Mg through phloem, and of Ca mainly through xylem,but only limited intake of Na and Cl. The next phase up to fullleaf expansion showed similar relative differences in xylemintake between ions and the onset of rapid phloem export fromthe lamina of K and Mg, some export of Na and Cl but scarcelyany of Ca. The next mature phase, marked by maximal photosynthesisand transpiration by the leaf, showed high xylem intake of allions in xylem. This was more than matched by phloem export ofMg and K, but by only fractional re-export of Na and Cl andagain very limited cycling through the leaf of Ca. The finalpre-senescence phase exhibited similar behaviour, but with generallygreater contribution to phloem transport from mobilization ofion reserves of the lamina. The petiole retained particularlylarge amounts of Na and Cl in its early growth, thereby protectingthe lamina from excessive entry of salt, but these petiolarpools, together with those or other nutrient ions, were laterpartially mobilized to the lamina via the xylem stream. Datawere discussed in relation to the relatively high salt toleranceexhibited by the species. Key words: Ricinus communis, xylem and phloem transport, ion balance, K+ economy, Na+ exclusion, NaCl-stress, salt tolerance, leaf development     

Fine Structure of Hydrolysed Primary Walls in Tracheary Elements of Petiolar Xylem in Eucalyptus delegatensis

The hydrolysed lateral primary walls of tracheary elements ofthe petiolar xylem of Eucalyptus delegatensis were examinedby electron microscopy. Vessel-vessel and vessel—tracheidhydrolysed walls were strikingly different in appearance fromtracheid—tracheid walls. The difference seemed to be inthe degree to which the primary walls were hydrolysed. The observationssuggest the wall hydrolysis to be an ordered and controlledprocess. Eucalyptus delegatensis, hydrolysed wall, petiolar xylem, tracheary elements     

Cation and Chloride Partitioning through Xylem and Phloem within the Whole Plant of Ricinus communis L. under Conditions of Salt Stress

Uptake and partitioning through the xylem and phloem of K⁺,Na⁺, Mg²⁺ , Ca²⁺ and Cl^– were studied over a 9 d intervalduring late vegetative growth of castor bean (Ricinus communisL.) plants exposed to a mean salinity stress of 128 mol m^–3NaCl. Empirically based models of flow and utilization of eachion within the whole plant were constructed using informationon ion increments of plant parts, molar ratios of ions to carbonin phloem sap sampled from petioles and stem internodes andpreviously derived information on carbon flow between plantsparts in xylem and phloem in identical plant material. Salientfeatures of the plant budget for K⁺ were prominent depositionin leaves, high mobility of K⁺ in phloem, high rates of cyclingthrough leaves and downward translocation of K⁺ providing theroot with a large excess of K⁺ . Corresponding data for Na⁺showed marked retention in the root, lateral uptake from xylemby hypocotyl, stem internodes and petioles leading to low intakeby young leaf laminae and substantial cycling from older leavesback to the root. The partitioning of the anionic componentof NaCl salinity, Cl^–, contrasted to that of Na⁺ in thatit was not substantially retained in the root, but depositedmore or less uniformly in stem, petiole and leaf lamina tissues.The flow pattern for Mg²⁺ showed relatively even depositionthrough the plant but some preferential uptake by young leaves,generally lesser export than import by leaf laminae, and a returnflow of Mg²⁺ from shoot to root considerably less than the recordedincrement of the root. Ca²⁺ partitioning contrasted with thatof the other ions in showing extremely poor phloem mobility,leading to progressive preferential accumulation in leaf laminaeand negligible cycling of the element through leaves or root.Features of the response of Ricinus to salinity shown in thepresent study were discussed with data from similar modellingstudies on white lupin (Lupinus albus L.) and barley (Hordeumvulgare L.) Key words: Ricinus communis L, potassium, sodium, chloride, calcium, magnesium, phloem, xylem, transport, partitioning, salinity     

Concentrations and Transport of Solutes in Xylem and Phloem along the Leaf Axis of NaCl-treated Hordeum vulgare

Hordeum vulgare cv. California Mariout was established in sandculture at two different NaCl concentrations (0.5 mol m^–3‘control’ and 100 mol m^–3) in the presenceof 6.5 mol m^–3 K ⁺. Between 16 and 31 d after germination,before stem elongation started, xylem sap was collected by useof a pressure chamber. Collections were made at three differentsites on leaves 1 and 3: at the base of the sheath, at the baseof the blade, i.e. above the ligule, and at the tip of the blade.Phloem sap was collected from leaf 3 at similar sites throughaphid stylets. The concentrations of K ⁺, Na⁺, Mg2⁺ and Ca²⁺were measured. Ion concentrations in xylem sap collected at the base of leaves1 and 3 were identical, indicating there was no preferentialdelivery of specific ions to older leaves. All ion concentrationsin the xylem decreased from the base of the leaf towards thetip; these gradients were remarkably steep for young leaves,indicating high rates of ion uptake from the xylem. The gradientsdecreased with leaf age, but did not disappear completely. In phloem sap, concentrations of K⁺ and total osmolality declinedslightly from the tip to the base of leaves of both controland salt-treated plants. By contrast, Na⁺ concentrations inphloem sap collected from salt-treated plants decreased drasticallyfrom 21 mol m^–3 at the tip to 7.5 mol m^–3 at thebase. Data of K/Na ratios in xylem and phloem sap were used to constructan empirical model of Na⁺ and K⁺ flows within xylem and phloemduring the life cycle of a leaf, indicating recirculation ofNa⁺ within the leaf. Key words: Hordeum vulgare, xylem transport, phloem transport, NaCl-stress     

The Effect of Reduced Hydraulic Conductance on Stomatal Conductance and Xylem Cavitation

The vulnerability of xylem conduits to cavitation theoreticallydetermines the maximum flow rate of water through plants, andhence maximum transpiration (E), stomatal conductance (g_s),and leaf area (A₁. Field-grown Betula occidentalis with a favourablewater supply exhibit midday xylem pressures (     

The Relationship between Endogenous Hormone Levels in the Plant and Seasonal Aspects of Cambial Activity

The auxin levels at different heights in a ring-porous (Ulmusglabra) and a diffuse-porous (Populus trichocarpa) tree werefollowed in spring. There was shown to be a correlation betweenthe pattern of spread of cambial activity and auxin gradients.The observed continuation of cambial activity after cessationof extension growth in ring-porous species is apparently dueto the fact that the mature leaves produce auxin, whereas littleauxin is produced by the leaves of diffuse-porous species underthese conditions. The spring/summerwood transition in ring-porousspecies can be related to a fall in auxin level at the timethat shoot extension-growth ceases. There is evidence of phloemproduction after the cessation of xylem production in Ailanthusaltissima and Robinia pseudacacia, but not in Betula pubescentand Acer pseudoplatanus. It is suggested that, in Ailan-thus,phloem production continues because high endogenous gibberellinlevels are present after the auxin level has been reduced byshort-day treatment.     

Co-ordinated development of the leaf midrib xylem with the lamina in Nicotiana tabacum

Background and Aims

The water-transport capacity of leaf venation is positively related to the leaf-lamina area, because the number and diameter of vein-xylem conduits are controlled to match the lamina area. This study aimed to investigate how this co-ordinated relationship between the leaf-lamina area and vein-xylem characteristics is achieved by examining the midrib xylem of tobacco leaves.

Methods

The changes in the midrib-xylem characteristics over time were quantified using leaves with four different final lamina areas. The measured data were fitted to sigmoidal functions. From the constants of the fitted curves, the final values in mature leaves, maximal developmental rates (V_Dev) and developmental duration (T_Dev) were estimated for each of the xylem characteristics. Whether it is the lamina or the midrib xylem that drives the co-ordinated development was examined by lamina removal from unfolding leaves. The effects of the application of 0·1 % IAA (indole-3-acetic acid) to leaves with the laminas removed were also analysed.

Key Results

For both the leaf lamina and the midrib-xylem characteristics, the differences in final values among leaves with different lamina areas were more strongly associated with those in V_Dev. Notably, the V_Dev values of the midrib-xylem characteristics were related to those of the leaf-lamina area. By lamina removal, the conduit diameter was reduced but the number of conduits did not significantly change. By IAA application, the decrease in the conduit diameter was halted, and the number of conduits in the midrib xylem increased.

Conclusions

According to the results, the V_Dev values of the lamina area and the midrib-xylem characteristics changed in a co-ordinated manner, so that the water-transport capacity of the midrib xylem was positively related to the leaf-lamina area. The results also suggest that IAA derived from the leaf lamina plays a crucial role in the development of the leaf venation.     

The Relationship Between Size of Shoot Apices and Morphological Features of Mature Leaves and Stems of Four Species of Angiosperm Trees

Shoot apex, leaf and stem growth parameters for four speciesof deciduous trees were measured. Only in elm was there a correlationbetween the size of shoot apical meristems and mature leaves.In ash, basswood and cottonwood there was no significant differencebetween size of shoot apices of sucker and canopy branches,despite significant differences in lamina size. In the suckerbranches of all species studied there occurred an early, lateralexpansion of the subapical region of the shoot apical meristem.This correlated well with the greater diameter of stem and pithregions of sucker branches. In addition, the season's annualring of xylem was greater in basswood, cottonwood and elm. Diametersof vessel elements were greater in sucker than canopy branchesin three of the four species. Total branch and internode andnumber of nodes per branch were significantly greater for suckerbranches than canopy growth of all species studied. A hypothesis is proposed to explain the development of the verylarge surface area of leaves on sucker branches. This hypothesisis based on the position of sucker branches in relation to theroot system and involves differences in water stress known tobe present in all plants.Copyright 1993, 1999 Academic Press Sucker leaves, canopy leaves, Fraxìnus pennsylvanica Marsh, green ash, Ulmus amerìcana L., American elm, Populus deltoides Marsh, cottonwood, Tilia americana, basswood     

On the Conductivity of the Xylem in Trees

The xylem of three tree species, the ring porous Fraxinus excelsiorL., and the diffuse porous Acer pseudoplatanus L., and Salixatrocinerea Brot., has been examined to determine how closelyflow through it obeys the Poiseuille equation. The equationwas most closly obeyed in the relationship between flow-rateand pressure difference between the ends of the xylem cylinder,though there were differences between the ring and diffuse poroustypes. The relationship of flow-rate and specific conductivityto the radius of the xylem cylinder was complex.Fraxinus showeda rise with increase of radius up to a given size (0.54cm),followed by a steep decline. This decline was shown to be causedby a progressive increase in the number of vessels which wereunable to conduct water. Small increases in the viscosity ofthe xylem liquid produced a disproportionately large fall inthe rate of flow, and the specific conductivity of the xylemdecreased. With viscosities greater than 2.5 centipoise, thespecific conductivity remained constant. An effect of alternatelyraising and lowering the hydrostatic pressure of the xylem wateron the specific conductivity of the wood is described. These results are discussed in relation to the structure ofthe xylem, and particularly with regard to the possibility thatthe conduction channels in the wood consist of a system of microcapillariessurrounding the vessel cavities.     

Water Flow Through Cotton Roots in Relation to Xylem Anatomy

The effect of root anatomy on water flow was studied in 7-d-oldcotton (Gossypium hirsutum L.) seedlings grown in solution culture.The total water flux of the intact root system was measuredusing a pressure chamber. Then successive terminal root sectionswere removed at 2,6,10 and 12 cm behind the root tip and theflux was remeasured after each successive cut was made. Xylemdevelopment at different distances behind the root apex wasstudied with a microscope using sections cut free-hand and stainedwith toluidine blue. Water flux increased with the removal ofsuccessive terminal root sections and this coincided with thedegree of basipetal primary xylem development. The large increasein water flux at 10 to 12 cm was associated with secondary xylemdevelopment and increased xylem vessel number. A comparison of water flow and xylem anatomy between roots withtetrarch (Stoneville 506 and Deltapine 41) and pentarch (T25strain) vascular bundle arrangements showed no significant differencesin the measured values of water flux for the primary root. Waterflux, estimated using Poiseuille's equation and measured xylemdimensions, was greater for the tetrarch roots, primarily becauseof the larger diameter of individual vessel elements. The increasednumber of vessel elements in the pentarch primary root of T25did not result in any apparent decrease in axial resistanceto water flow. Key words: Gossypium hirsutum L., roots, vascular bundle, xylem water flux, xylem     

Haustorium-related Uptake and Metabolism of Host Xylem Solutes by the Root Hemiparasitic ShrubSantalum acuminatum(R. Br.) A. DC. (Santalaceae)

Solute composition of root xylem sap of common native hostsof quandong (Santalum acuminatum) was compared with that ofcorresponding xylem sap and ethanolic extracts of endophytictissues of haustoria of the hemiparasite. Each host transporteda characteristic set of organic nitrogenous solutes, but littleor no nitrate, and the data indicated only limited direct flowof amino compounds between xylem streams of hosts and parasite.Proline predominated in the haustorium and xylem ofSantalum,but was at negligible levels in the xylem of most hosts. Sucrose,fructose, glucose, malate and citrate were at high levels inall saps, and fructose especially prominent inSantalum. Chloride,sulphate and phosphate were the principal inorganic anions ofthe xylem. Based on C:N ratios of xylem and dry matter ofSantalumandassuming a 70% or more dependence on the host for N, it wasestimated thatSantalumwould gain approximately one third ofits C requirement for dry matter production heterotrophicallyfrom the xylem of its hosts. Infiltration of xylem of haustoria-bearingroot segments of a major host (Acacia rostellifera) with a rangeof¹⁵N labelled substrates resulted in 40–80% of the¹⁵Nof endophytes of the attached haustoria being received as proline.Nitrate reductase activity was induced in haustoria followinghost xylem feeding of nitrate. The study concludes that haustoriaofSantalumact as a major site of synthesis and export of prolineand might therefore play an important role in osmotic adjustmentof the parasite and its related acquisition of water from hosts. Root hemiparasite; Santalum acuminatum; ¹⁵N labelled substrates; xylem transport; proline; osmoregulation     

Antitranspirant Activity in Xylem Sap of Maize Plants

Xylem sap from unwatered maize plants was collected and testedfor antitranspirant activity. Two assays were used. These werea transpiration assay with detached wheat leaves and a stomatalbio-assay involving the direct microscopic observation of epidermisof Commelina communis. The reduction in transpiration of detached wheat leaves promotedby xylem sap could be duplicated almost exactly by the applicationof solutions of ABA of equivalent concentration to that foundin the xylem sap. Removal of virtually all the ABA from thexylem sap, using an immunoaffinity column, removed virtuallyall the antitranspirant activity in both assays. These results are discussed in the context of other resultswhich suggest the presence of as-yet unidentified inhibitorsin the xylem sap of unwatered plants. We suggest that with maize plants at least, stomatal responsesto soil drying can be entirely explained by enhanced concentrationof ABA in the xylem stream. Key words: Antitranspirant activity, ABA, ABA bio-assay, xylem sap     

Xylem Sap from Actinidia chinensis: Gradients in Sap Composition

In early spring, just prior to leaf break, there were substantialgradients in the concentration of many solutes in xylem sapcollected by vacuum extraction from shoots of different ages,from the branches, trunk and roots of mature Chinese gooseberryvines (Actinidia chinensis Planchon) For most nutrients therewas a progressive fall in concentration from youngest shootsdown to the roots and nitrate was the only nutrient to showa trend in the opposite direction These gradients in concentrationare probably sufficient to account for most of the changes observedin the composition of bleeding xylem sap during the first fewhours of bleeding after a branch is cut or wounded Actinidia chinensis, Chinese gooseberry, kiwifruit, xylem sap, nutrient gradients     

Membrane Potentials in the Xylem in Roots of Intact Plants

The membrane potential differences (PDs) of root cells of intact,illuminated Trifolium repens L. and Lolium perenne L. have beenmeasured. In T. repens the PDs were the same for all cell typesexcept for the xylem vessels, which were more positive, andfor some cells immediately adjacent to the xylem vessels whichwere 10 mV more negative. The mean PD for all cells was emdash164.6 ± 0.6 mV and the mean for cells adjacent to thexylem vessels with elevated PDs was 178.4 ± 2.4 mV. Whenthe electrode tip was in a xylem vessel a low but stable PD(mean = emdash 89.9 mV) was recorded. The results for L. perennewere similar except that there were no cells with elevated PDsadjacent to the xylem vessels. An inhibitor of ion transport from the root to the shoot, p-fluorophenylalanine(p-FPA), caused a depolarization of 10 mV in the cell PDs butin the xylem vessels the depolarization was 50 mV. The possibility that the elevated PDs of cells adjacent to thexylem vessels are related to the transport of ions into thevessels is discussed.     

Adaptive Growth Responses to Osmotic Stress of Hypocotyl Sections of Vigna unguiculata: Roles of the Xylem Proton Pump and IAA

The growth responses to osmotic stress of hypocotyl sectionsof Vigna unguiculata were studied by the xylem perfusion method.Hypocotyl sections shrank upon exposure to osmotic stress. Sectionsshowed no adaptive responses to osmotic stress when they werein an IAA-depleted condition as a result of perfusion with solutionsthat lacked IAA for 3–4 h. The correlation between thegrowth rate and the membrane potential of the xylem/symplastboundary (V^px) was very limited in the absence of IAA. By contrast,hypocotyl sections showed distinct adaptive responses to osmoticstress after perfusion with solutions that contained 10 µMIAA. In the presence of IAA, V^px increased in the negative directionand growth resumed in spite of the osmotic stress. The growthrate was closely correlated with the xylem membrane potential.Hyper-polarization of the membranes of the xylem/symplast boundaryalways preceded the recovery of growth under osmotic stress.It appears that IAA is essential for the adaptive recovery ofgrowth under osmotic stress and, moreover, that the xylem protonpump plays an indispensable role in modulating the growth ofhypocotyl sections. This result confirms prediction of an earliersimulation study using the apoplast canal model [Katou and Furumoto(1986) Protoplasma 133: 174, Katou and Enomoto (1991) PlantCell Physiol. 32: 343]. (Received June 27, 1996; Accepted October 28, 1996)     

The Characteristics of the Adjustment in the pH of the Xylem Exudate of Segments of Vigna Hypocotyls during Xylem Perfusion

The xylem vessels of segments of Vigna hypocotyl were perfusedwith solutions of various pH values. With both acid and alkalineperfusion solutions, the pH of the xylem exudate tended to equilibrateat approximately 6 during the course of xylem perfusion. Thisphenomenon was observed both in air and under anoxia, and alsoin segments pretreated with methanol or preheated in a microwaveoven. Therefore, the adjustment in pH does not depend on therespiratory metabolism but originates in the buffering actionof the xylem wall apoplast. High concentrations of protons inducedthe release of other cations into the xylem exudate under anoxia,and high concentrations of K⁺ or Ca²⁺ ions induced the releaseof protons into the xylem under anoxia. These results indicatethat a cation-exchange reaction on the xylem cell wall was responsiblefor the buffering effect. The physiological role of the highbuffering capacity of the xylem wall apoplast is discussed inlight of the role of the proton pump of the plasmalemma in elongationgrowth. (Received September 25, 1990; Accepted January 21, 1991)     

Three Different Methods for Measuring Xylem Cavitation and Embolism: A Comparison

Three different methods for measuring xylem embolism due towater cavitation were compared—the acoustic method, thehydraulic method and the anatomical method. Young plants ofCeratonia siliqua L. were water stressed for 9, 16 and 23 d. Xylem cavitation was detected by counting ultrasound (100–300kHz) acoustic emissions (AE) from 1-year-old twigs (acousticmethod). Xylem embolism was detected by measuring the loss ofhydraulic conductivity of twigs of the same age (hydraulic method).The blockage of single xylem conduits was detected by perfusingSafranin into the xylem of 1-year-old twigs of stressed plantsand measuring the number and the diameters of non-conductingxylem conduits, under the microscope (anatomical method). It was noted that: (a) the number of AE and the loss of conductivityincreased with the water stress applied; (b) a linear relationseemed to exist between the number of AE and the loss of conductivity,suggesting that the AE counted could be only (or mainly) producedin the xylem conduits; (c) the vulnerability of the xylem conduitsto embolism was a direct function of their diameter; and (d)the measured loss of conductivity was of the same order of magnitudeas the theoretical one. The three methods gave fairly similar results. Nonetheless,they are not alternative to one another in that: (a) the acousticmethod allows continuous recordings to be made but does notprovide information about the actual damage suffered by plants;(b) the hydraulic method is very informative but destructive;and (c) the anatomical method is very useful both in phytogeographicaland in genetic improvement studies. Ceratonia siliqua L, Carob tree, water stress, xylem embolism, acoustic method, hydraulic method, anatomical method