Vacuolar Na/K Exchange, its Occurrence in Root Cells of Hordeum, Atriplex and Zea and its Significance for K/Na Discrimination in Roots

Using excised low-salt roots of barley and Atriplex hortenslsthe transport of endogenous potassium through the xylem vesselswas studied It was enhanced by nitrate and additionally by sodiumions which apparently replaced vacuolar potassium which wasthen available in the symplasm of root cells for transport tothe shoot Vacuolar Na/K exchange also has been investigatedby measurements of longitudinal ion profiles in single rootsof both species. In Atriplex roots a change in the externalsolution from K⁺ to Na⁺ induced an exchange of vacuolar K⁺ forNa⁺, in particular in the subapical root tissues and led toincreased K⁺ transport and loss of K⁺ from the cortex. In inverseexperiments a change from Na⁺ to K⁺ did not induce an exchangeof vacuolar Na⁺; merely in meristematic tissues Na⁺—apparentlyfrom the cytoplasm—was extruded in exchange for K⁺. Inroots of barley seedlings without caryopsis, as in excised roots,a massive exchange of K⁺ for Na⁺ was observed in the continuouspresence of external 1.0 mM Na and 0.2 mM K. This exchange alsowas attributed to the vacuole and was most pronounced in theyoung subapical tissues. It did not occur, however, in the correspondingtissues in roots of fully intact barley seedlings. In these,the young tissues retained a relatively high K/Na ratio alsoin their vacuoles. Similarly, contrasting results were obtainedwith intact and excised roots of Zea mays L. Based on theseresults a scheme of the events that lead to selective cationuptake in intact barley roots is proposed. In this scheme acrucial factor of selectivity is sufficient phloem recirculationof K⁺ by the aid of which K⁺ rich cortical cells are formednear the root tip. When matured these cells are suggested tomaintain a high cytoplasmic K/Na ratio due to K⁺ dependent sodiumextrusion at the plasmalemma and due to recovery of vacuolarK⁺ by Na/K exchange across the tonoplast. Key words: Potassium/Sodium selectivity, Vacuolar exchange, Xylem transport, Hordeum, Zea, Atriplex     

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     

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     

Determinants of vertebrate invasion success in Europe and North America

Species that are frequently introduced to an exotic range have a high potential of becoming invasive. Besides propagule pressure, however, no other generally strong determinant of invasion success is known. Although evidence has accumulated that human affiliates (domesticates, pets, human commensals) also have high invasion success, existing studies do not distinguish whether this success can be completely explained by or is partly independent of propagule pressure. Here, we analyze both factors independently, propagule pressure and human affiliation. We also consider a third factor directly related to humans, hunting, and 17 traits on each species' population size and extent, diet, body size, and life history. Our dataset includes all 2362 freshwater fish, mammals, and birds native to Europe or North America. In contrast to most previous studies, we look at the complete invasion process consisting of (1) introduction, (2) establishment, and (3) spread. In this way, we not only consider which of the introduced species became invasive but also which species were introduced. Of the 20 factors tested, propagule pressure and human affiliation were the two strongest determinants of invasion success across all taxa and steps. This was true for multivariate analyses that account for intercorrelations among variables as well as univariate analyses, suggesting that human affiliation influenced invasion success independently of propagule pressure. Some factors affected the different steps of the invasion process antagonistically. For example, game species were much more likely to be introduced to an exotic continent than nonhunted species but tended to be less likely to establish themselves and spread. Such antagonistic effects show the importance of considering the complete invasion process.     

Potassium Fluxes in Excised Barley Roots

The method of the modified compartmental analysis for excisedroots has been adopted for measuring K⁺-fluxes and compartmentationin barley (Hordeum distichon) roots. Efflux of ⁴²K and ⁸⁶Rbindicated that more than two intracellular compartments wereinvolved in the tracer exchange; the ⁴²K data clearly showedthe components. On the basis of the efflux behaviour of theapical and more basal tissues of the roots, the three componentsof efflux were attributed to the cytoplasm of differentiated(fast) and meristematic tissues (intermediate) and to the vacuoles(slow exchange) of the roots. A model is proposed on the basisof which, the fluxes corresponding to the meristematic and differentiatedtissues of the root can be estimated. Additionally, fluxes ofthe differentiated root tissues were determined by using effluxdata obtained with root segments without apical tissues. Thedata obtained in both ways compare reasonably well and agreeto independent chemical measurements. Comparison of the ⁴²K and ⁸⁶Rb efflux data show strong discriminationof K^– in favour of Rb⁺ and indicate that ⁸⁶Rb is not suitableas a tracer for K⁺ in efflux measurements, at least with barleyroots.