Idioblast Cells Research Articles

Secretory Structure, Histochemistry and Phytochemistry Analyses of Stimulant Plant

Plants that are used as stimulant supposed to contains various metabolit compounds that are produced or secreted by secretory structures. This study aimed to identify the secretory structure of plant used as stimulant and chemical compounds accumulated in it. The secretory structure and its histochemistry were observed on plant material that are used as herbal ingredient. Phytochemical content was analyzed by using a qualitative test. The result showed that the idioblast cells and secretory cavities were found in the leaves of Decaspermum fruticosum, and Polyalthia rumphii. Most idioblast cells contained lipophilic substances and terpenoids or alkaloids, while secretory cavity contained alkaloid. Phytochemical analysis for D. fruticosum, and P. rumphii contain terpenoids, phenols, steroids, and flavonoids

IDENTIFICATION OF SECRETORY STRUCTURE, HISTOCHEMISTRY AND PHYTOCHEMICAL COMPOUNDS OF MEDICINAL PLANT Hyptis capitata Jacq.

Hyptis capitata Jacq. (common name: Knobweed or False Ironwort) belongs to Lamiaceae family and is among known traditional medicine. The Anak Dalam Tribe of Jambi Province uses the leaves of H. capitata to cure external and internal wounds. This study was aimed at identifying and analyzing secretory structure, histochemistry and phytochemical content of H. capitata leaves.  The results showed that H. capitata leaves have secretory structures in the form of peltate, capitate and uniseriate glandular trichomes on the upper and lower leaf surfaces, with idioblast cells scattered throughout the leaf mesophyll. Histochemical tests indicated that the peltate trichomes have four head cells, containing alkaloids and terpenoids. This study classified capitate trichomes into two types: 1. type I has one stalk cell and two head cells, all containing  alkaloids, terpenoids and lipophilic compounds; 2. type II has long stalk cells consisting of seven to ten cells with one head cell, all containing alkaloids and terpenoids. Uniseriate glandular trichomes of H. capitata leaves consist of four to eight cells containing alkaloids and terpenoids. The idioblast cells are round-shaped and contain lipophilic compounds. GC-MS analysis showed that H. capitata leaves  contain terpenoid compounds assumed to serve as anti-infective agents, including l-limonene, eugenol, farnesol isomers A, d-nerolidol, hexahydrofarnesol and neophytadiene.

Cell-specific localization of alkaloids in Catharanthus roseus stem tissue measured with Imaging MS and Single-cell MS

Catharanthus roseus (L.) G. Don is a medicinal plant well known for producing antitumor drugs such as vinblastine and vincristine, which are classified as terpenoid indole alkaloids (TIAs). The TIA metabolic pathway in C. roseus has been extensively studied. However, the localization of TIA intermediates at the cellular level has not been demonstrated directly. In the present study, the metabolic pathway of TIA in C. roseus was studied with two forefront metabolomic techniques, that is, Imaging mass spectrometry (MS) and live Single-cell MS, to elucidate cell-specific TIA localization in the stem tissue. Imaging MS indicated that most TIAs localize in the idioblast and laticifer cells, which emit blue fluorescence under UV excitation. Single-cell MS was applied to four different kinds of cells [idioblast (specialized parenchyma cell), laticifer, parenchyma, and epidermal cells] in the stem longitudinal section. Principal component analysis of Imaging MS and Single-cell MS spectra of these cells showed that similar alkaloids accumulate in both idioblast cell and laticifer cell. From MS/MS analysis of Single-cell MS spectra, catharanthine, ajmalicine, and strictosidine were found in both cell types in C. roseus stem tissue, where serpentine was also accumulated. Based on these data, we discuss the significance of TIA synthesis and accumulation in the idioblast and laticifer cells of C. roseus stem tissue.

Comparative effects of water deficit and enhanced UV-B radiation on photosynthetic capacity and leaf anatomy traits of two grapevine (Vitis vinifera L.) cultivars

The effects of drought and enhanced UV-B radiation on photosynthetic and leaf anatomical characteristics were studied in two different grapevine genotypes (Vitis vinifera L. cvs ‘Romeiko’ and ‘Soultanina’), which were grown outdoors in 25 L pots containing a peat:perlite:sand (3:1:1, v/v/v) potting mix. The plants were subjected to three levels of UV-B radiation (i) ambient, (ii) intermediate (ambient plus 15 % UV-B radiation), (iii) high (ambient plus 30 % UV-B radiation) and two irrigation regimes (i) well-watered (irrigated on daily basis to substrate capacity) and (ii) drought-stressed (irrigated daily with the 50 % of the amount of water provided to well-watered plants). Both abiotic stressors induced a significant decrease in net photosynthesis (Pn) while the combination of drought and elevated UV-B radiation appeared to have synergistic effects on photosynthetic rate. At leaf anatomical level, drought resulted in the development of idioblastic cells in leaves of both grapevine varieties as well as in the reduction of leaf expansion rate, total lamina thickness and total biomass per leaf area whereas leaf tissue density considerably increased. On the other hand, enhanced UV-B radiation did not affect leaf anatomical characteristics of both genotypes in a similar way. In particular, different responses, closely depended to UV-B levels applied, were observed concerning the chlorenchymatic tissue and total lamina thickness. Moreover, the stimulation of UV-B absorbing compounds biosynthesis under enhanced UV-B radiation, could not manage to preserve photosynthetic performance. The implication of the watering regime and the UV-B in the induction of leaf anatomical and morphological alterations in net photosynthesis is discussed.

Myrosin idioblast cell fate and development are regulated by the Arabidopsis transcription factor FAMA, the auxin pathway, and vesicular trafficking.

Crucifer shoots harbor a glucosinolate-myrosinase system that defends against insect predation. Arabidopsis thaliana myrosinase (thioglucoside glucohydrolase [TGG]) accumulates in stomata and in myrosin idioblasts (MIs). This work reports that the basic helix-loop-helix transcription factor FAMA that is key to stomatal development is also expressed in MIs. The loss of FAMA function abolishes MI fate as well as the expression of the myrosinase genes TGG1 and TGG2. MI cells have previously been reported to be located in the phloem. Instead, we found that MIs arise from the ground meristem rather than provascular tissues and thus are not homologous with phloem. Moreover, MI patterning and morphogenesis are abnormal when the function of the ARF-GEF gene GNOM is lost as well as when auxin efflux and vesicular trafficking are chemically disrupted. Stomata and MI cells constitute part of a wider system that reduces plant predation, the so-called "mustard oil bomb," in which vacuole breakage in cells harboring myrosinase and glucosinolate yields a brew toxic to many animals, especially insects. This identification of the gene that confers the fate of MIs, as well as stomata, might facilitate the development of strategies for engineering crops to mitigate predation.

Idioblastic mucilage cells in Teucrium polium leaf. Anatomy and histochemistry

In the mesophyll of Teucrium polium L. leaves, isolated or grouped idioblastic secretory cells occur in contact with the distal (adaxial) vessel elements of the vascular bundles. They appear to originate from one or more bundle sheath cells and their intracellular space is entirely occupied by the secretory material. The latter has a glycoproteinaceous constitution (mucilage), as histochemical tests showed. Idioblastic cells, therefore, correspond to typical mucilage cells. Mucilage seems to play a crucial role in the adaptation of the plant to unfavourable environmental conditions.

Chemical composition and ruminal nutrient degradability of fresh and ensiled amaranth forage

Amaranth is a crop with potential as a source of forage for ruminants that has not been well characterized. A study was conducted to determine the impact of ensiling on the nutritional quality and ruminal degradability of forage from two amaranth cultivars adapted to North America (i.e. Plainsman and D136). In particular, quantification and some microscopic characterization of oxalate found in amaranth were performed as it is an antiquality compound of concern. There were limited interactions between cultivars and ensiling for most variables. Differences in chemical composition between amaranth cultivars were also limited. Ensiling reduced non-structural carbohydrate and true protein contents. The proportion of acid detergent protein was high in fresh and ensiled forages of both cultivars (average of 177 g kg(-1) crude protein). Total oxalate content averaged 30 and 25 g kg(-1) in fresh and ensiled forages respectively. Ensiling reduced soluble oxalate content. Crystals observed in amaranth were calcium oxalate druses found mostly in idioblast cells in leaf mesophyll and parenchyma of primary and secondary veins. In situ ruminal degradability data indicated that both fresh and ensiled amaranth are highly degradable in the rumen. This study confirms that amaranth is a suitable forage for ruminant animals. Its chemical composition is comparable, for most variables, to that of other commonly used forage species.

English

A fundamental understanding of mango flowering in the tropics is essential to efficiently utilize cropping management systems which could extend both the flowering and crop production seasons. However, the information and appreciation of the floral biology of this popular fruit species is still lacking. Therefore, the objective of this work was to observe the morphology of Chok Anan mango flowers using scanning electron microscope. The Chok Anan mango flower is monoecius where both the male and hermaphrodite flowers exist in the same panicle. The fruit develops from the hermaphrodite flower while the male flower contributes the pollen when pollination occurred. The Chok Anan mango flowers contained five sepals and five petals arranged in a whorl. There were not much differences in the structure of sepals and petals between the male and hermaphrodite flowers. The structure of the petals is not flat but billowy. Both sepals and petals consist of ground parenchyma tissue with laticifer and idioblast cells, starch granules and vascular bundle tissues. Trichomes are present in both sepals and petals. The male flower has the same structure as the hermaphrodite flower except that it does not have carpel. Further study is needed to understand the peculiarities of the floral morphology to allow us to predict mango production. Key words: Pollen, sepal, petal, hermaphrodite, floral biology.

Heterogeneity of silica and glycan-epitope distribution in epidermal idioblast cell walls in Adiantum raddianum laminae

Laminae of Adiantum raddianum Presl., a fern belonging to the family Pteridaceae, are characterised by the presence of epidermal fibre-like cells under the vascular bundles. These cells were thought to contain silica bodies, but their thickened walls leave no space for intracellular silica suggesting it may actually be deposited within their walls. Using advanced electron microscopy in conjunction with energy dispersive X-ray microanalysis we showed the presence of silica in the cell walls of the fibre-like idioblasts. However, it was specifically localised to the outer layers of the periclinal wall facing the leaf surface, with the thick secondary wall being devoid of silica. Immunocytochemical experiments were performed to ascertain the respective localisation of silica deposition and glycan polymers. Epitopes characteristic for pectic homogalacturonan and the hemicelluloses xyloglucan and mannan were detected in most epidermal walls, including the silica-rich cell wall layers. The monoclonal antibody, LM6, raised against pectic arabinan, labelled the silica-rich primary wall of the epidermal fibre-like cells and the guard cell walls, which were also shown to contain silica. We hypothesise that the silicified outer wall layers of the epidermal fibre-like cells support the lamina during cell expansion prior to secondary wall formation. This implies that silicification does not impede cell elongation. Although our results suggest that pectic arabinan may be implicated in silica deposition, further detailed analyses are needed to confirm this. The combinatorial approach presented here, which allows correlative screening and in situ localisation of silicon and cell wall polysaccharide distribution, shows great potential for future studies.

Fatty acid profile and elemental content of avocado (Persea americana Mill.) oil –effect of extraction methods

Interest in vegetable oil extracted from idioblast cells of avocado fruit is growing. In this study, five extraction methods to produce avocado oil have been compared: traditional solvent extraction using a Soxhlet or ultrasound, Soxhlet extraction combined with microwave or ultra-turrax treatment and supercritical fluid extraction (SFE). Traditional Soxhlet extraction produced the most reproducible results, 64.76 ± 0.24 g oil/100 g dry weight (DW) and 63.67 ± 0.20 g oil/100 g DW for Hass and Fuerte varieties, respectively. Microwave extraction gave the highest yield of oil (69.94%) from the Hass variety. Oils from microwave extraction had the highest fatty acid content; oils from SFE had wider range of fatty acids. Oils from Fuerte variety had a higher monounsaturated: saturated FA ratio (3.45–3.70). SFE and microwave extraction produced the best quality oil, better than traditional Soxhlet extraction, with the least amount of oxidizing metals present.

‘Chambourcin’ grapevine (Vitis vinifera Vitis L.) berries display sustained post-veraison accumulation of oxalic acid

SUMMARYA substantial accumulation of organic acids (OAs) occurs during pre-veraison grape berry development, which results in low pH conditions and contributes to the organoleptic properties of the fruit. The main OAs in most grape berries are tartaric acid (TA) and malic acid (MA). MA is accumulated via a primary metabolic shunt, whereas TA is accumulated via ascorbic acid (AA; vitamin C) metabolism. Oxalic acid (OX) is also derived from AA in competition with TA, and results from a cleavage between the second and third carbon atoms. Oxalic acid is subsequently biomineralised as calcium oxalate crystals in idioblast cells. We have examined the biosynthesis of TA, MA, and OX during grape berry development in four grapevine cultivars. We show that in the Vitis vinifera L. Vitis L. complex inter-specific hybrid, ‘Chambourcin’, there is a high capacity to accumulate OX during post-veraison fruit development. OX content increased from approx. 1 mg g–1 to 3 mg g–1 berry fresh weight between 14 and 16 weeks post-anthesis, resulting in OX levels that were similar to the MA levels in mature fruit. Whether OX accumulation occurs due to increased biomineralisation or biosynthesis, our results show that ‘Chambourcin’ grape berries represent the only documented case of post-veraison accumulation of OX.

Cell-specific expression of tryptophan decarboxylase and 10-hydroxygeraniol oxidoreductase, key genes involved in camptothecin biosynthesis in Camptotheca acuminata Decne (Nyssaceae).

BackgroundCamptotheca acuminata is a major natural source of the terpenoid indole alkaloid camptothecin (CPT). At present, little is known about the cellular distribution of the biosynthesis of CPT, which would be useful knowledge for developing new strategies and technologies for improving alkaloid production.ResultsThe pattern of CPT accumulation was compared with the expression pattern of some genes involved in CPT biosynthesis in C. acuminata [i.e., Ca-TDC1 and Ca-TDC2 (encoding for tryptophan decarboxylase) and Ca-HGO (encoding for 10-hydroxygeraniol oxidoreductase)]. Both CPT accumulation and gene expression were investigated in plants at different degrees of development and in plantlets subjected to drought-stress. In all organs, CPT accumulation was detected in epidermal idioblasts, in some glandular trichomes, and in groups of idioblast cells localized in parenchyma tissues. Drought-stress caused an increase in CPT accumulation and in the number of glandular trichomes containing CPT, whereas no increase in epidermal or parenchymatous idioblasts was observed. In the leaf, Ca-TDC1 expression was detected in some epidermal cells and in groups of mesophyll cells but not in glandular trichomes; in the stem, it was observed in parenchyma cells of the vascular tissue; in the root, no expression was detected. Ca-TDC2 expression was observed exclusively in leaves of plantlets subjected to drought-stress, in the same sites described for Ca-TDC1. In the leaf, Ca-HGO was detected in all chlorenchyma cells; in the stem, it was observed in the same sites described for Ca-TDC1; in the root, no expression was detected.ConclusionsThe finding that the sites of CPT accumulation are not consistently the same as those in which the studied genes are expressed demonstrates an organ-to-organ and cell-to-cell translocation of CPT or its precursors.

ANATOMICAL STUDIES ON WILD SOLANUM TAXA IN EGYPT

The comparative study on anatomical structure of stems and leaves in 12 wild Solanum taxa re-vealed that the outline of cross sections of stem and petiole, presence or absence of hypodermis, structure of cortex, pericycle and types of steles vary considerably in different studied taxa. It was also evident that other anatomical features of leaf including mesophyll types, arrangement and num-ber of the bundles at the mid-rib region, presence or absence of bundle sheath and idioblast cells as well as the vascular system of petiole are important diagnostic characters in the proposed key for iden-tification. A key including variations in anatomical characters of stems and leaves of the investigated taxa is provided

Morpho-anatomical features of the leaves of the mediterranean geophyteUrginea maritima (L) baker (Liliaceae)

The morpho-anatomy and histochemistry of the hysteranthous leaf ofUrginea maritima (L.) Baker and its adaptive strategies to the Mediterranean climate were investigated. The leaf ofU. maritima is 714 μm thick and possesses moderate specific leaf mass (8.564 mg cm-2) and low tissue density (136.5 mg cm-3). The epidermal cells are compactly arranged and covered with cuticle. The average density of stomata in lower epidermis is higher than that of the upper one. The mesophyll cells occupy 52.96% of the total volume of the leaf, while the mesophyll intercellular spaces and the air spaces occupy 30.41%. Idioblastic cells containing raphide bundles and different phenotypes of crystalloid inclusions, embedded in polysaccharides, occur in the lower side of the mesophyll. The presence of oil droplets and lipids is evident. Bundle sheath cells are hardly visible with no chloroplasts which are a pronounced C3 plant character. Plastids containing protein crystalloid inclusions are abundant in the protophloem sieve elements.U. maritima, a deciduous plant, possesses leaves with mesophytic characters, in order to optimize its adaptation to the seasonal fluctuation of environmental conditions of the Mediterranean climate.

Rhythmic plant morphogenesis: Recurrent patterns of idioblast cell production

Most plants are constructed from repeating modular units such as phytomers, merophytes, and cell packets. Even an organism as simple as the filamentous cyanobacterium Anabaena shows recurrent patterns of differentiated cellular structures, notably with respect to its heterocysts. These examples reflect the inherent rhythms established within developmental processes of living organisms. In the present article, attention is paid to repetitious production of idioblasts—isolated cells, or clusters of cells, with an identity different to that of neighbouring cells from which they are derived. In higher plant root tissues, idioblasts are contained within cell packets that grow up from mother cells during the course of a number of cycles of cell production. The heterocysts of Anabaena are also discussed; they, too, are a type of idioblast. The idioblasts of root tissues originate as small cells which result from unequal cell divisions. Such divisions are usually the final ones within a cell packet which has already undergone a number of division cycles and are characteristically located at one or both ends of a packet. The packet end walls are suggested to have a role in regulating division asymmetry. Idioblastic systems discussed are root cortical trichosclereids and diaphragm cells; in their earliest stage, the cells from which lateral root primordia arise are also considered as clusters of idioblasts because they, too, are the products of asymmetric divisions of pericyclic mother cells. The division patterns of all these idioblastic systems were modelled in a consistent way using L-systems, with the assumption that the age of a cell-packet end wall plays a special role in cell determination.

Calcium carbonate deposition in a cell wall sac formed in mulberry idioblasts

Although calcium carbonate is known to be a common biomineral in plants, very little attention has been given to the biological control of calcium carbonate deposition. In mulberry leaves, a subcellular structure is involved in mineral deposition and is described here by a variety of cytological techniques. Calcium carbonate was deposited in large, rounded idioblast cells located in the upper epidermal layer of mulberry leaves. Next to the outmost region ("cap") of young idioblasts, we found that the inner cell wall layer expanded to form a peculiar outgrowth, named cell wall sac in this report. This sac grew and eventually occupied the entire apoplastic space of the idioblast. Inside the mature cell wall sac, various cellulosic membranes developed and became the major site of Ca carbonate deposition. Concentrated Ca2+ was pooled in the peripheral zone, where small Ca carbonate globules were present in large numbers. Large globules were tightly packed among multiple membranes in the central zone, especially in compartments formed by cellulosic membranes and in their neighboring membranes. The maximum Ca sink capacity of a single cell wall sac was quantified using enzymatically isolated idioblasts as approximately 48 ng. The newly formed outgrowth in idioblasts is not a pure calcareous body but a complex cell wall structure filled with substantial amounts of Ca carbonate crystals.

Seed coat morphology of Aizoaceae–Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance

Macro- and micromorphological characters of the seeds of 32 species belonging to Aizoaceae (26 species), Gisekiaceae (one species) and Molluginaceae (five species) were analysed for their taxonomic value. Seed morphology is found to be of considerable taxonomic value within the taxa investigated, although additional criteria are usually necessary for distinguishing the species. However, a subgroup of Trianthema, consisting of T. cussackiana, T. megasperma, T. pilosa, T. rhynchocalyptra and T. oxycalyptra var. oxycalyptra (Aizoaceae) shares the synapomorphy of seeds with scattered idioblast cells (papillae). There is little infrageneric variation within Sesuvium and Zaleya in contrast to Trianthema, whose seeds are considerably polymorphic. The fairly different seed structures found in Aizoaceae s.s. and Molluginaceae support their treatment as two distinct families. However, the position of Lineum (Molluginaceae) remains uncertain. Moreover, seed characters support the independence of Gisekia (Gisekiaceae) in a family of its own.

Processes modulating calcium distribution in citrus leaves. An investigation using x-ray microanalysis with strontium as a tracer.

Citrus leaves accumulate large amounts of calcium that must be compartmented effectively to prevent stomatal closure by extracellular Ca2+ and interference with Ca(2+)-based cell signaling pathways. Using x-ray microanalysis, the distribution of calcium between vacuoles in different cell types of leaves of rough lemon (Citrus jambhiri Lush.) was investigated. Calcium was accumulated principally in palisade, spongy mesophyll, and crystal-containing idioblast cells. It was low in epidermal and bundle sheath cells. Potassium showed the reverse distribution. Rubidium and strontium were used as tracers to examine the pathways by which potassium and calcium reached these cells. Comparisons of strontium and calcium distribution indicated that strontium is a good tracer for calcium, but rubidium did not mirror the potassium distribution pattern. The amount of strontium accumulated was highest in palisade cells, lowest in bundle sheath and epidermal cells, and intermediate in the spongy mesophyll. Accumulation of strontium in palisade and spongy mesophyll was accompanied by loss of potassium from these cells and its accumulation in the bundle sheath. Strontium moved apoplastically from the xylem to all cell types, and manipulation of water loss from the adaxial leaf surface suggested that diffusion is responsible for strontium movement to this side of the leaf. The results highlight the importance of palisade and spongy mesophyll as repositories for calcium and suggest that calcium distribution between different cell types is the result of differential rates of uptake. This tracer technique can provide important information about the ion uptake and accumulation properties of cells in intact leaves.

Wood and stem anatomy of woody Amaranthaceae s.s.: ecology, systematics and the problems of defining rays in dicotyledons

Wood and stem anatomy is studied for seven species of six genera (root anatomy also reported for one species) of Amaranthaceae s.s. Quantitative data on vessels correlate closely with relative xeromorphy of respective species, agreeing with values reported for dicotyledons without successive cambia in comparable habitats. Libriform fibre abundance increases and vessel diameter decreases as stems and roots of the annual Amaranthus caudatus mature. Long, thick-walled fibres in Bosea yervamora may be related to the upright nature of elongate semi-climbing stems. Non-bordered or minutely bordered perforation plates characterize Amaranthaceae, as they do most other Caryophyllales. Amaranthaceae have idioblastic cells containing druses, rhomboidal crystals or crystal sand: these forms intergrade and seem closely related. Rays are present in secondary xylem of the Amaranthaceae studied. Cells intermediate between ray cells and libriform fibres occur in Charpentiera elliptica. Degrees of diversity in rays and reports of raylessness in Amaranthaceae induce discussion of definition and identification of rays in dicotyledons; some sources recognize both rays and radial plates of conjunctive tissue in Amaranthaceae. The action of successive cambia is described: lateral meristem periclinal divisions produce secondary cortex externally, conjunctive tissue internally and yield vascular cambia as well. Vascular cambia produce secondary phloem and secondary xylem, in both ray and fascicular zones, as in a dicotyledon with a single cambium. Identification of meristem activity and appreciation of varied ray manifestations are essential in understanding the ontogeny of stems in Amaranthaceae (which have recently been united with Chenopodiaceae). © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143, 1–19.

Calcium-accumulating cells in the meristematic region of grapevine root apices.

Apical roots of grapevines were examined by cryo-SEM (scanning electron microscopy) and the intracellular distribution of Ca was demonstrated by X-ray microanalysis in different regions of the primary root. We show that large amounts of Ca are accumulated as raphide crystals in the vacuoles of specialised cortical cells (idioblast cells) of the root apex. These crystal idioblast cells appeared to form a discontinuous cone of cells in the outer region of the root meristem. The raphide crystals within these cells were less apparent in older regions of the root, 10-12 mm basipetal to the root tip. We suggest that the raphide crystals could initially act as another Ca sink involved in the regulation of Ca levels in root apices. In older regions of the root these cells are spaced at intervals around the periphery of the cortex and the subsequent disappearance of the raphides may be indicative of remobilisation, perhaps in the zone of elongation where cell wall synthesis occurs and Ca demand is high. Calcium-accumulating cells were also observed in the older regions of the root, forming endodermal protrusions extending into the cortex. These cells may play a part in regulating Ca delivery to the xylem stream by sequestration of Ca from the radial flow of water at the endodermis. The observed distribution of Ca in root apices was different from the other major cations (e.g. K) and anions (e.g. Cl) because high concentrations were localised to specific cells. We interpret the results in the context of a model of the dynamics of grapevine root growth and cell differentiation, and the temporal balance of solute supply from the protophloem and the external medium.