Leaf-like Structure in the Photosynthetic, Succulent Stems of Cacti

This research examined the hypothesis that as cacti evolve tothe leafless condition, the stem epidermis and cortex becomemore leaflike and more compatible with a photosynthetic role.All cacti in the relict genus Pereskia have non-succulent stemsand broad, thin leaves. All members of the derived subfamilyCactoideae are ‘leafless’, having an expanded cortexthat is the plant's only photosynthetic tissue. In Pereskia,leaves have a high stomatal density (mean: 50.7 stomata mm^–2in the lower epidermis, 38.1 mm^–2 in the upper epidermis),but stems have low stomatal densities (mean: 11.3 mm ₂, threeof the species have none). Stems of Cactoideae have a high stomataldensity (mean: 31.1 mm^–2, all species have stomata). Theouter cortex cells of stems of Cactoideae occur in columns,forming a palisade cortex similar to a leaf palisade parenchyma.In this palisade cortex, the fraction of tissue volume availablefor gas diffusion has a mean volume of 12.9%, which is identicalto that of Pereskia leaf palisade parenchyma. Pereskia stemcortex is much less aerenchymatous (mean: 5.3% of cortex volume).Cactoideae palisade cortex has a high internal surface density(0.0207 cm₂ cm^–2 which is higher than in Pereskia stemcortex (0.0150 cm₂ cm^–3) but not as high as Pereskia leafpalisade parenchyma (0.0396 cm₂ cm^–3). Pereskia stem cortexhas no cortical bundles, but Cactoideae cortexes have extensivenetworks of collateral vascular bundles that resemble leaf veins. Cactaceae, cactus, intercellular space, stomatal density, internal surface/volume, evolution     

Time Course and Polarity of Primary Phloem Fibre Differentiation in Pisum sativum

Primary phloem fibres of Pisum sativum deposit lignified cellwalls, 2 and 5 days after germination in root and stem, respectively.Fibre bundles reach their final size within 4–6 days.The differentiation of the bundles as a whole is faster in thestem compared with the root. In the special diverging bundlesof the lower internodes of the stem the peripheral portionsmature earlier than the inner portions. A wound in the rootas well as in the stem, interrrupts the differentiation of fibresdirectly below it. At least one bundle below a wound is disturbedand often a whole bundle is missing. The meaning of these findingsconcerning the control of cell size is discussed. Pisum sativum, differentiation, induction, phloem fibres, polarity, time course     

The Composition of Phloem Exudate and Xylem Sap from Tree Tobacco (Nicotiana glauca Grah.)

The composition of xylem sap and exudate from stem incisionsof Nicotiana glauca Grah. was compared in detail. Exudationfrom stem incisions occurred over a 5 min period in certainplants, enabling collection of 5–30 µl of sap. Therate of exudation showed an exponential decline. Exudate hada high dry matter content (170–196 mg ml^–1) andhigh sugar (sucrose) levels. Xylem sap had a low pH (5.8) andexudate a pH of 7.9. Glutamine dominated the amino compoundsin xylem sap and exudate, and K⁺ was the major cation. Totalamino compounds in stem exudate reached 10.8 mg ml^–1 whereasxylem sap contained much lower levels (0.28 mg ml^–1).All mineral elements and amino compounds with the exceptionof calcium were more concentrated in stem exudate than in xylemsap. Sucrose was labelled heavily in stem exudate following pulsingof an adjacent leaf with ¹⁴CO₂. A concentration gradient ofsugar (2.1 bar m^–1) was recorded for stems. Levels ofsucrose, amino compounds and K⁺ ions in stem exudate showeda diurnal periodicity. Each commodity reached maximum concentrationat or near noon and minimum concentration about dawn. The evidencesuggests that exudate from stem incisions of N. glauca is arepresentative sample of solutes translocated in the phloem. Nicotiana glauca Grah., phloem sap, xylem sap, sucrose, amino compounds, mineral ions     

VASCULAR ORGANIZATION IN SHOOTS OF CACTACEAE. I. DEVELOPMENT AND MORPHOLOGY OF PRIMARY VASCULATURE IN PERESKIOIDEAE AND OPUNTIOIDEAE

Primary shoot vasculature has been studied for 31 species of Pereskioideae and Opuntioideae from serial transections and stained, decorticated shoot tips. The eustele of all species is interpreted as consisting of sympodia, one for each orthostichy. A sympodium is composed of a vertically continuous axial bundle from which arise leaf- and areole-trace bundles and, in many species, accessory bundles and bridges between axial bundles. Provascular strands for leaf traces and axial bundles are initiated acropetally and continuously within the residual meristem, but differentiation of procambium for areole traces and bridges is delayed until primordia form on axillary buds. The differentiation patterns of primary phloem and xylem are those typically found in other dicotyledons. In all species vascular supply for a leaf is principally derived from only one procambial bundle that arises from axial bundles, whereas traces from two axial bundles supply the axillary bud. Two structural patterns of primary vasculature are found in the species examined. In four species of Pereskia that possess the least specialized wood in the stem, primary vascular systems are open, and leaf traces are mostly multipartite, arising from one axial bundle. In other Pereskioideae and Opuntioideae the vascular systems are closed through a bridge at each node that arises near the base of each leaf, and leaf traces are generally bipartite or single. Vascular systems in Pereskiopsis are relatively simple as compared to the complex vasculature of Opuntia, in which a vascular network is formed at each node by fusion of two sympodia and a leaf trace with areole traces and numerous accessory bundles. Variations in nodal structure correlate well with differences in external shoot morphology. Previous reports that cacti have typical 2-trace, unilacunar nodal structure are probably incorrect. Pereskioideae and Opuntioideae have no additional medullary or cortical systems.     

Regeneration of Tissues in Wounded Stems: A Quantitative Study

When a dicotyledonous stem is wounded by longitudinally splittinga young internode into halves, cells near the cut surface proliferateto form a callus within which vascular tissues differentiateand tend to restore a vascular cylinder in each half. Threephases of regeneration after wounding were identified and quantifiedin stems of three Solanaceous species. (1) In an initial ‘lag’phase, lasting about 2 d, neither cell division nor enlargementwere detected, but mitotic figures were observed within about300 µm of the cut surface. (2) Throughout a second, ‘division’phase, from about days 2–10, cell division and enlargementoccurred. Both were initiated mainly in the two cell layersnearest the surface. A mass of callus formed, with new cellwalls mostly parallel to the surface. Cell enlargement laggedbehind cell division for the first few days, so that mean radialcell diameter decreased until day 6, thereafter remaining almostconstant at 30–40 µm. Towards the end of this phase,mitoses ceased within the callus except in the positions ofthe future vascular and cork cambia, where radial cell diameterfell towards a constant 15–20 µm. (3) During a third,‘differentiation’ phase, cell division was restrictedto the cambial zones, and derivatives differentiated into cork,phloem or xylem according to position. The rate of increasein cell number per transect was 1.5–2.0 cells d^–1,of which more than half was xylem. Capsicum annuum L., sweet pepper, Lycopersicon esculentum Mill., tomato, cambium, cell division, differentiation, regeneration, wounding of stems, xylem     

Quantum Flux Density as a Factor Controlling the Rate of Growth, Carbohydrate Partitioning and Wood Structure of Betula pubescens Seedlings

The ability of Betula pubescens seedlings to acclimate to arange of quantum flux densities is examined, paying particularattention to the quantity, rate and type of wood produced. Betulapubescens seedlings were germinated and grown for 2 months ata quantum flux density of 440 µmol m^–2 s^–1,then transferred to 440, 244 or 89 µmol m^–2 s^–1. Seedlings transferred to reduced fluence rates were able tosurvive and grow. Acclimation was associated with an increasedallocation of carbon to stem elongation and leaf-area production,this occurred at the expense of root storage carbohydrate androot growth. The rate of wood production was shown to be directlyrelated to quantum flux density. Stem wood structure at smallquantum flux density showed an increase in proportion of fibres,while root wood structure in this regime showed a greater proportionof vessels. In general the activity of phenylalanine ammonia-lyase(PAL) showed little significant difference between treatments.The implications of these changes in wood structure are discussedand related to wood production and PAL activity. Betula pubescens Ehrh., birch., shade tolerance, carbohydrate partitioning, relative growth rate, wood structure, phenylalanine-ammonia-lyase     

Xylem and Phloem Transport of Mineral Nutrients from Solanum tuberosum Roots

The xylem and phloem transport of mineral elements from stemnodal roots to the stem and stolon of growing potato (Solanumtuberosum L. cv. ‘Russet Burbank’) plants was investigated.Adventitious roots, originating from below-ground nodes of thestem of potato seedlings, were exposed to solutions of SrCI₂or MnSO₄. Relative elemental concentrations were measured inthe conductive tissues using energy dispersive X-ray analysis.After a 5 h daylight uptake period, Sr (a Ca-transport analogue)levels were elevated in the stem xylem tissue, but Sr did notincrease in the stem phloem, nor was it present in either ofthe conductive tissues of stolons located 1–2 nodes abovethe treated roots. In contrast, elevated levels of Cl, S, andMn were found in stolon xylem and phloem tissue during the sameperiod. The absence of Sr in the stolon after 5 h suggests thatno xylem flow into the stolon occurred during the uptake periodand, furthermore, phloem flow is responsible for the transportof the Cl, S, and Mn into the stolon. Elevated levels of thesemobile nutrients in the xylem of the stolon were attributedto xylem-to-phloem transfer in the stem or leaves, transportto the stolon in the phloem, and phloem-to-xylem transfer inthe stolon. During a 19 h uptake period, some Sr was observedin the phloem tissue of the stem, demonstrating slow exchangeof Sr with sieve elements or proximal phloem parenchyma andcompanion cells. Key words: Calcium, manganese, X-ray analysis     

Water-storing and Cavitation-preventing Adaptations in Wood of Cacti

Ancestral cacti presumably had abundant, fibrous, heavily lignifiedwood, similar to that present in the relictual, leaf-bearinggenus Pereskia. During the evolutionary radiation of the subfamilyCactoideae, diverse types of bodies and woods arose. Severalevolutionary lines have retained an abundant, fibrous wood:all wood cells, even ray cells, have thick lignified walls,and axial parenchyma is only scanty paratracheal. Aside froma diversity of vessel diameters, there seems to be little protectionagainst cavitation during water-stress, and little water-storagecapacity. This strong wood permits the plants to be tall andto compete for light in their tree-shaded semi-arid habitats.In other evolutionary lines, the wood lacks fibres, and almostall cells have thin, unlignified walls. Vessels occur in anextensive matrix of water-storing parenchyma, and tracheidsare also abundant, constituting over half the axial tissue insome species. There is excellent protection against cavitation,but little mechanical support for the plant body; however, theseplants are short and occur in extremely arid, unshaded sites.Scandent, vinelike plants of two genera produce a dimorphicwood—while their shoots are extending without externalsupport, they produce fibrous, lignified wood, but after leaningagainst a host branch, they produce a parenchymatous, unlignifiedwood.Copyright 1993, 1999 Academic Press Cactaceae, cactus, water-stress, wood, evolution, xylem     

Boron Mobility and Nutrition in Broccoli (Brassica oleracea var. italica

Broccoli (Brassica oleracea var. italica cv Premium Crop) plantswere germinated in soil, transferred to vermiculite three weekslater and grown in the glasshouse, then either supplied continuouslywith boron levels ranging from 0.0 (deficient) to 12.5 (toxic)mg l^–1 of nutrient solution or transferred from 2.5 to0.0 mg B l^–1 at the initiation of inflorescence development.At commercial maturity the concentrations of various inorganicand organic solutes in phloem exudates and xylem saps, as wellas plant characteristics and elemental composition of the variousplant parts, were determined. Under deficient B levels leaf midrib and stem corkiness wereevident, together with signs of stem pith breakdown, symptomswhich resemble the initiation of the hollow stem disorder. Thexylem sap B concentration declined by about 50 % when B wasnot supplied or was removed after a period of adequate supply;the phloem concentration was unaffected. Also, the decreasingB concentration gradients from mature transpiring tissues toyoung developing sinks disappeared. Therefore, it is concludedthat when B is deficient, it is retranslocated from source leavesin the phloem stream supplying the developing leaves and inflorescence.The data also suggested that at toxic levels B undergoes extensivelateral transfer, probably from xylem to xylem, thereby enhancingthe B concentration of developing sinks. The B regime influenced dry-matter partitioning, retranslocationof some elements, and the synthesis and distribution of aminoacids and sugars, reflecting the general nature of B involvementin plant processes. Brassica oleracea var., italica, broccoli, phloem, mobility, retranslocation, boron nutrition, transport fluids, concentration gradients     

Growth of Zea mays L. plants with their seminal roots only. Effects on plant development, xylem transport, mineral nutrition and the flow and distribution of abscisic acid (ABA) as a possible shoot to root signal

Maize (Zea mays L.) was grown in quartz sand culture eitherwith a normal root system (controls) or with seminal roots only(‘single-rooted’). Development of adventitious rootswas prevented by using plants with an etiolated mesocotyl andthe stem base was positioned 5–8 cm above the sand. Eventhough the roots of the single-rooted plants were sufficientlysupplied with water and nutrients, the leaves experienced waterdeficits and showed decreased transpiration as trans plrationalwater flow was restricted by the constant number of xylem vesselspresent in the mesocotyl. As a consequence of this restriction,transpirational water flow velocities in the metaxylem vesselsreached mean values of 270 m h^–1 and phloem transportvelocities of 5.2 m h^–1. Despite limited xylem transportmineral nutrient concentrations in leaf tissues were not decreasedin single-rooted plants, but shoot and particularly stem developmentwas somewhat inhibited. Due to the lack of adventitious rootsthe shoot:root ratio was strongly increased in the single-rootedplants, but the seminal roots showed compensatory growth comparedto those in control plants. Consistent with decreased leaf conductance,ABA concentrations in leaves of single-rooted plants were elevatedup to 10-fold, but xylem sap ABA concentrations in these plantswere lower than in controls, in good agreement with the well-wateredconditions experienced by the seminal roots. Surprisingly, however,ABA concentrations in tissues of the seminal roots of the single-rooted plants were clearly increased compared to the controls,presumably due to increased ABA import via phloem from the water-stressedleaves. The results are discussed in relation to the role ofABA as a shoot to root signal. Key words: Zea mays, seminal roots, plant development, xylem transport, mineral nutrition, ABA, shoot-to-root signal     

How to become a tree without wood – biomechanical analysis of the stem of Carica papaya L

Carica papaya L. does not contain wood, according to the botanical definition of wood as lignified secondary xylem. Despite its parenchymatous secondary xylem, these plants are able to grow up to 10‐m high. This is surprising, as wooden structural elements are the ubiquitous strategy for supporting height growth in plants. Proposed possible alternative principles to explain the compensation for lack of wood in C. papaya are turgor pressure of the parenchyma, lignified phloem fibres in the bark, or a combination of the two. Interestingly, lignified tissue comprises only 5–8% of the entire stem mass. Furthermore, the phloem fibres do not form a compact tube enclosing the xylem, but instead form a mesh tubular structure. To investigate the mechanism of papaya's unusually high mechanical strength, a set of mechanical measurements were undertaken on whole stems and tissue sections of secondary phloem and xylem. The structural Young's modulus of mature stems reached 2.5 GPa. Since this is low compared to woody plants, the flexural rigidity of papaya stem construction may mainly be based on a higher second moment of inertia. Additionally, stem turgor pressure was determined indirectly by immersing specimens in sucrose solutions of different osmolalities, followed by mechanical tests; turgor pressure was between 0.82 and 1.25 MPa, indicating that turgor is essential for flexural rigidity of the entire stem.     

两种寄生植物营养器官结构的研究

研究了槲寄生和北桑寄生茎、叶和吸器的结构,结果表明：槲寄生三年生茎稍肉质化,无周皮,有极厚的角质层;皮层有7个纤维束,薄壁组织发达;木质部中亦含有大量薄壁组织;韧皮部发达;簇晶及大量粘稠物质在茎中广泛分布。槲寄生叶肉质化,厚1.44 mm; 角质层发达,厚度为9.06～13.4 μm;叶肉无栅栏组织和海绵组织的分化。北桑寄生茎与正常木本植物茎的结构相似,木质部为典型的环孔材;韧皮部含有石细胞;髓部细胞壁厚,单纹孔明显。北桑寄生叶较薄,叶脉发达,无栅栏组织和海绵组织的分化。两种植物的吸器结构相似,由薄壁组织和螺纹导管组成,它们的螺纹导管长度相同,但槲寄生导管的直径是北桑寄生的2.5倍。     

Vascular Continuity in the Primary and Secondary Stem Tissues of Bougainvillea (Nyctaginaceae)

An investigation of stem structure of Bougainvillea by serialsections and cine-photography shows that the medullary systemof the inner area of young stems is the sole vascular systemdirectly continuous into the lateral appendages (leaves, axillarybuds and axillary thorns) via complex nodal anastomoses. Thevascular system at the periphery of the primary bundles is notdirectly continuous into these appendages. In secondary growth,there is direct continuity between vascular bundles within asingle ring, in a tangential direction via either xylem aloneor both xylem and phloem, and between rings in a radial directionalways via xylem and phloem, even though the rings are derivativesof successive cambia. Bougainvillea, vascular system, phloem, xylem, anomalous secondary thickening     

Anatomical Adaptations to Xeric Conditions in Maihuenia (Cactaceae), a Relictual, Leaf-Bearing Cactus

Maihuenia and Pereskia, constitute Pereskioideae, the subfamily of Cactaceae with the greatest number of relictual features, but the two genera differ strongly in habit and ecological adaptations. Plants of Maihuenia occur in extremely xeric regions of Patagonia and are small cushion plants with reduced, terete leaves and soft, slightly succulent trunks. Plants of Pereskia occur only in mesic or slightly arid regions and are leafy trees with hard, woody trunks and thin, broad leaves. Maihuenias have many more anatomical adaptations to arid conditions than do pereskias: maihuenias lack sclerenchyma in their phloem and cortex (M. poeppigii also lacks xylem sclerenchyma and can contract during drought); their wood consists of vessels, axial parenchyma, and wide-band tracheids and can store water as well as minimize embolism damage; one species channelizes water flow by producing intraxylary bark; and at least some stem-based photosynthesis occurs because maihuenias have small patches of persistent stem epidermis that bears stomata and overlies a small amount of aerenchymatous chlorenchyma. Pereskias lack all these features. Although closely related, maihuenias have fewer relictual features than do pereskias, and plants of Pereskia probably are more similar to the ancestral cacti. Received 8 March 1999/ Accepted in revised form 29 May 1999     

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     

Transport of Cadmium via Xylem and Phloem in Maturing Wheat Shoots: Comparison with the Translocation of Zinc, Strontium and Rubidium

The toxic heavy metal cadmium is taken up by plants and maycontaminate harvested parts of agricultural crops. In the experimentsreported here, cadmium was introduced together with markersfor phloem (rubidium) and xylem (strontium) transport, eitherinto intact shoots via a flap below the flag leaf node, or intodetached shoots via the cut stem. Cadmium introduced into intactplants was redistributed during maturation from the peduncleand the flag leaf lamina to the grain. In detached shoots, somecadmium was removed from the transpiration stream, as judgedfrom the comparison of shoots steam-girdled below the ear andof controls with an intact phloem in the peduncle. A minor quantityof cadmium was transported to the grain via the phloem in controlshoots while a high percentage of this element was retainedin the peduncle. The cadmium content of the grain increasedin response to the increased cadmium concentrations in the feedingsolutions (0.1 to 10 µM). The cadmium content of the grainwas slightly lower when zinc (>10 µM) was introducedat the same time as cadmium (1 µM).Copyright 1997 Annalsof Botany Company Triticum aestivumL.; cadmium; phloem transport; wheat; zinc     

Xylem fluxes of fixed N through nodules of the legume Acacia littorea and haustoria of an associated N-dependent root hemiparasite Olax phyllanthi

Nodulated 1-1.5-year-old plants of Acacia littorea grown inminus nitrogen culture were each partnered with a single seedlingof the root hemiparasite Olax phyllanthi. Partitioning of fixedN between plant organs of the host and parasite was studiedfor the period 4–8 months after introducing the parasite.N fluxes through nodules of Acacia and xylem-tapping haustoriaof Olax were compared using measured xylem flows of fixed Nand anatomical information for the two organs. N₂ fixation duringthe study interval (635 µg N g FW nodules^–1 d^–1)corresponded to a xylem loading flux of 0.20 µg N mm^–2d^–1 across the secretory membranes of the pencycle parenchymaof the nodule vascular strands. A much higher flux of N (4891µg mm^–2 d^–1) exited through xylem at the junctionof nodule and root. The corresponding flux of N from host xylemacross absorptive membranes of the endophyte parenchyma of Olaxhaustorium was 1.15 µg N mm^–1 d^–1, six timesthe loading flux in nodules. The exit flux from haustorium toparasite rootlet was 20.0 pg N mm^–1 d^–1, 200-foldless than that passing through xylem elements of the nodule.Fluxes of individual amino compounds in xylem of nodule andhaustorium were assessed on a molar and N basis. N flux valuesare related to data for transpiration and partitioning of Cand N of the association recorded in a companion paper. Key words: Olax phyllanthi, host-parasite relationships, N flux, Acacia, N₂ fixation     

Modelling of the Partitioning, Assimilation and Storage of Nitrate within Root and Shoot Organs of Castor Bean (Ricinus communis L.)

An experimentally-based modelling technique was developed todescribe quantitatively the uptake, flow, storage and utilizationof NO₃-N over a 9 d period in mid-vegetative growth of sandcultured castor bean (Ricinus communis L.) fed 12 mol m^–3nitrate and exposed to a mean salinity stress of 128 mol m^–3NaCl. Model construction used information on increments or lossesof NO₃-N or total reduced N in plant parts over the study periodand concentration data for NO₃-N and reduced (amino acid) Nin phloem sap and pressure-induced xylem exudates obtained fromstem, petiole and leaf lamina tissue at various levels up ashoot. The resulting models indicated that the bulk (87%) of incomingnitrate was reduced, 51% of this in the root, the remainderprincipally in the laminae of leaves. The shoot was 60% autotrophicfor N through its own nitrate assimilation, but was oversuppliedwith surplus reduced N generated by the root and fed to theshoot through the xylem. The equivalent of over half (53%) ofthis N returned to the root as phloem translocate and, mostly,then cycled back to the shoot via xylem. Nitrate comprised almosthalf of the N of most xylem samples, but less than 1% of phloemsap N. Laminae of leaves of different age varied greatly inN balance. The fully grown lower three leaves generated a surplusof reduced N by nitrate assimilation and this, accompanied byreduced N cycling by xylem to phloem exchange, was exportedfrom the leaf. Leaf 4 was gauged to be just self-sufficientin terms of nitrate reduction, while also cycling reduced N.The three upper leaves (5–7) met their N balance to varyingextents by xylem import, phloem import (leaves 6 and 7 only)and assimilation of nitrate. Petioles and stem tissue generallyshowed low reductase activities, but obtained most of theirN by abstraction from xylem and phloem streams. The models predictedthat nodal tissue of lower parts of the stem abstracted reducedN from the departing leaf traces and transferred this, but notnitrate, to xylem streams passing further up the shoot. As aresult, xylem sap was predicted to become more concentratedin N as it passed up the shoot, and to decrease the ratio ofNO₃-N to reduced N from 0·45 to 0·21 from thebase to the top of the shoot. These changes were reflected inthe measured N values for pressure-induced xylem exudates fromdifferent sites on the shoot. Transfer cells, observed in thexylem of leaf traces exiting from nodal tissue, were suggestedto be involved in the abstraction process. Key words: Ricinus communis, nitrogen, nitrate, nitrate reduction, partitioning, phloem, xylem, flow models     

Some Experimental and Theoretical Observations Concerning Mass Flow in the Vascular Bundles of Heracleum

The theory and practice of applying the thermodynamics of irreversibleprocesses to mass-flow theories is presented. Onsager coefficientswere measured on cut and uncut phloem and cut xylem strandsof Heracleum muntegazzimum. In 0.3 N sucrose + 1 mN KC1 theyare as follows. In phloem, L_EE = 5 ? 10–⁴ mho cm^–1,L_pE = 9 ? 10^–6 cm³ s^–1 cm^–2 volt^–1 cm,and L_PP = 0.16 cm³ s^–1 cm^–2 (J cm^–3)^–1cm. In uncut phloem strands L_EE is about 1 ? 10^–3 mhocm^–1. In xylem in 2 x 10^–3 N KCI, L_pp = 50 to 225,L_PE = 2 ? 10^–4, and L^EE = 4 ? 10^–3. The measurementsare tentative since the blockage of the sieve plates is an interferingfactor, but if they are valid they lead to the conclusion thatneither a pressure-flow nor an electro-kinetic mechanism envisaginga ‘long distance’ current pathway can be the majormotive ‘force’ for transport in mature phloem. Measurementsof biopotentials along conducting but laterally detached phloembundles of Heracleum suggest, nevertheless, that there may bea small electro-osmotic component of at least 0.1 mV cm^–1endogenous in the phloem.