Richard Storey

Quaternary ammonium compounds in plants in relation to salt resistance

Abstract Fourteen plant species exhibiting a wide range of salt resistance as halophytes, semi-resistant glycophytes and sensitive glycophytes, have been grown in nutrient solution culture under low and high salt conditions. Inorganic analyses and shoot sap osmotic pressure values of these plants confirm that osmotic compensation at high salt levels is largely achieved by the accumulation of Na salts. Choline was found in shoots and roots in the range 1.0-0.2 μmol g fr. wt −1 and varied little following salt stress. Trigonelline was found in some of the sensitive glycophytes and did not increase significantly in stressed plants. Betaine levels were high (10 μmol g fr. wt −1 ) in the shoot of the halophytes at low salt conditions, lower values (1–10 μmol g fr. wt −1 ) were found in the semi-resistant glycophytes and none detected in the sensitive glycophytes. In the two resistant groups betaine accumulated to higher levels following NaCl stress. Shoot betaine levels always exceeded root levels. Proline occurred in all plants and in all cases was accumulated following NaCl stress.

Taxonomic and ecological aspects of the distribution of glycinebetaine and related compounds in plants

SummaryThe concentrations of the major inorganic ions and glycinebetaine, choline and proline and the osmotic pressure of extract sap have been determined in eight salt marsh species and four sand dune species from local habitats. These results together with those previously reported on hydroponically grown plants and data assembled from the literature show that glycinebetaine accumulation is a feature of members of the Chenopodiaceae, Amaranthaceae, many Gramineae and some members of the Solanaceae and Compositae, particularly when exposed to conditions of low soil water potential. It is suggested that in these families betaine is employed as a non-toxic cytoplasmic osmoticum when decreased osmotic potentials are required. In some other plant species proline may fulfil a similar function. Another quaternary ammonium compound may be accumulated in the Plumbaginaceae in addition to proline. Some evidence suggests that the differences in the organic osmoticum used may relate to the different inorganic ion contents of the plants. The accumulation of nitrogen dipoles as cytoplasmic osmotica may make heavy demands on the nitrogen economy of the plants and this problem is discussed briefly.

Betaine and choline levels in plants and their relationship to NaCl stress

Abstract The nature and concentration of quaternary ammonium compounds in 7 plants representing a spectrum of salt sensitivities under unstressed and NaCl-stressed conditions has been studied. The basal betaine levels and their elevation under stress suggest a correlation with the salt resistance of the plants. The shoot betaine levels in Rhodes grass at four different salt levels showed an approximately linear response to salt stress. The choline levels in all species were relatively constant and not correlated with salt resistance. As previously reported, the proline levels showed large increases in the glycophytes under stress but the response was somewhat different to that of betaine. The possible role of betaine in salt resistance is discussed.

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 Ca2+-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.

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.

Overexpression of a gibberellin inactivation gene alters seed development, KNOX gene expression, and plant development in Arabidopsis.

We have examined the role of gibberellins (GAs) in plant development by expression of the pea GA 2-oxidase2 (PsGA2ox2) cDNA, which encodes a GA inactivating enzyme, under the control of the MEDEA (MEA) promoter. Expression of MEA:PsGA2ox2 in Arabidopsis caused seed abortion, demonstrating that active GAs in the endosperm are essential for normal seed development. MEA:PsGA2ox2 plants had reduced ovule number per ovary and exhibited defects in phyllotaxy and leaf morphology which were partly suppressed by GA treatment. The leaf architecture and phyllotaxy defects of MEA:PsGA2ox2 plants were also restored by sly1-d which reduces DELLA protein stability to increase GA response. MEA:PsGA2ox2 seedlings had increased expression of the KNOTTED1-like homeobox (KNOX) genes, BP, KNAT2 and KNAT6, which are known to control plant architecture. The expression of KNOX genes is also altered in wild-type plants treated with GA. These results support the conclusion that GAs can suppress the effects of elevated KNOX gene expression, and raise the possibility that localized changes in GA levels caused by PsGA2ox2 alter the expression of KNOX genes to modify plant architecture.

Distribution of Potassium between Vacuole and Cytoplasm in Response to Potassium Deficiency

Differential changes in the distribution of potassium between cytoplasm and vacuole may be an important response of plants to K+ deficiency. Leigh and Wyn-Jones (1984) suggested that as tissue K+ concentration declines, the K+ concentration in the cytoplasm will remain constant while that in the vacuole will decrease. They proposed that decreases in cytoplasmic K+ concentration would only be observed at low tissue K+ concentrations and that growth would decline in response to these decreases. Evidence for such differential behaviour of the cytoplasmic and vacuolar K+ pools in barley leaf cells has now been obtained using X-ray microanalysis.