Photosynthesis of six barley genotypes as affected by water stress

The effect of water stress on plant water status and net photosynthetic gas exchange (PN) in six barley genotypes (Hordeum vulgare L.) differing in productivity and drought tolerance was studied in a controlled growth chamber. Osmotic adjustment (OA), PN, stomatal conductance (gs), and the ratio intercellular/ambient. CO2 concentration (Ci/Ca) were evaluated at four different levels of soil water availability, corresponding to 75, 35, 25 and 15 % of total available water. Variability in OA capacity was observed between genotypes: the drought tolerant genotypes Albacete and Alpha showed higher OA than drought susceptible genotypes Express and Mogador. The genotype Albacete exhibited also higher PN than the others at low water potential (Ψ). The ratios of PN/gs and Ci/Ca showed that differences in photosynthetic inhibition between genotypes at low Ψ were probably due to nonstomatal effects. In Tichedrett, a landrace genotype with a very extensive root development, OA was not observed, however, it exhibited a capacity to maintain its photosynthetic activity under water stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Pathogenesis of barley leaves by Helminthosporium teres II: Carbohydrate involvement

Conidia of Helminthosporium teres had negligible difference in germination and germ tube length between the decolorized and non-decolorized host leaves. Appressoria, penetration and colonization were less on decolorized host leaves, but addition of exogenous nutrients stimulated these stages. Leached conidia had reduced germination on decolorized host leaves, while appressoria formation, penetration and colonization were negligible. The addition of nutrients in the external environment, however, enabled some of the leached conidia to penetrate and colonize. Stimulation by the exogenous nutrients in decreasing order were: sucrose > glucose > yeast extract > leaf leachates. Optimum levels for various nutrients tested were 2% (w/v) each of sucrose and glucose, and 0.1% (w/v) yeast extract. Higher concentrations inhibited these stages of infection. Leached conidia and decolorized leaves had smaller amounts of carbohydrates than non-leached conidia and non-decolorized leaves, respectively. Depletion of host carbohydrates reduced appressoria formation, penetration and colonization and loss of carbohydrates from spores reduced germination.     

Weed–barley interactions affect plant acceptance by aphids in laboratory and field experiments

Increased botanical diversity can lead to suppression of insect pests. One route by which botanical diversity is increased in crops is through the occurrence of weeds, which increasingly interact with crop plants as organic production expands. However, the mechanisms by which this might affect insect herbivores are poorly understood. This study examined whether volatile chemical interactions between weeds and barley, Hordeum vulgare L. (Poaceae), can affect plant acceptance by the bird cherry oat aphid, Rhopalosiphum padi L. (Hemiptera: Aphididae). In laboratory experiments, exposure of barley to volatiles from Chenopodium album L. (Amaranthaceae) and Solanum nigrum L. (Solanaceae) resulted in significantly reduced aphid acceptance compared with unexposed plants. In a series of field experiments in which the occurrence of weeds was manipulated in plots of barley, significantly lower aphid acceptance was recorded on barley plants grown in plots with C. album compared with barley plants in weedless plots. The results indicate that interaction between weeds and barley can affect aphid–plant interactions in the field as well as in the laboratory and provide further evidence that the effects of chemical interactions between visibly undamaged plants can extend to higher trophic levels.     

Hormonal Regulation of Assimilate Utilization in Barley Leaves in Relation to the Development of Their Source Function

Growth, photosynthesis, utilization of assimilates, and the development of a source function in leaves were studied in relation to changes in concentrations and ratios of phytohormones. Carbon isotope ¹⁴C was used to trace utilization and outflow of photosynthetic products from the leaf. Concentrations of endogenous phytohormones were determined by solid-phase immunoenzyme assay. It was shown that, in juvenile leaves (one-fifth of their final area), which did not attain a high rate of photosynthesis, up to 80% of assimilates were incorporated into structural polysaccharides (cellulose and hemicellulose) one day after feeding with ¹⁴CO₂. During leaf growth and the development of its source function, the synthesis of structural polysaccharides declined to 10%, but the formation of alcohol- and water-soluble compounds (AWSC) grew to 80%. Monosaccharides and oligosaccharides, which could act as transport forms of carbohydrates, constituted 30% and 40% of AWSC, respectively. The percentage of assimilates utilized for protein synthesis decreased with leaf growth. The revealed changes correlate with the concentration and the ratio of free forms of phytohormones at various stages of leaf development. Development of a source function, a decline in cellulose and hemicellulose syntheses, and an increase in AWSC were related to the decrease in ABA and IAA concentrations and the increase in the ABA/IAA ratio. The ABA level can regulate the pathways of photoassimilate utilization in leaves by partitioning carbon flows either to the synthesis of high-molecular-weight compounds (cellulose, hemicellulose, and proteins), used for cell growth in leaves, or to the synthesis of transport forms of carbohydrates. A high ABA level favors the first pathway while low level switches leaf metabolism to the second one.     

Effect of calcium on sulfate uptake by barley roots

Abstract Sulfate uptake by excised roots of barley (Hordeum vulgare L.) was maximal in the presence of about 3x10^-3M CaCl₂. Kinetic studies contraindicate a stoichiometric binding of calcium to the carrier for sulfate, in contrast to findings of Cuppoletti and Segel (Biochemistry 14: 471–4718, 1975) for the filamentous fungus Penicillium notatum. In barley, calcium affects the K_m but not the V_max for sulfate uptake, presumably by altering the conformation and, thereby, the affinity of the carrier. Calcium also affects the transition site for sulfate uptake.     

Mannitol transport and suitability as an osmoticum in root cells

Fluxes of mannitol across plasmalemma and tonoplast of excised carrot storage root tissue were measured using compartmental analysis of ¹⁴C tracer exchange. Mannitol metabolism and the contribution of [¹⁴C]-labelled metabolites to efflux was shown to be small. Similar but less extensive measurements were made on red beet ( Beta vulgaris L.), barley ( Hordeum vulgare L.) and maize ( Zea mays L.) roots. Calculated values of the reflection coefficient for mannitol were close to one, but, despite this, the inflow of mannitol was sufficient to dissipate the mannitol concentration gradient between inside and outside the cells within the time it takes them to adjust vacuolar concentrations. Thus mannitol is not suitable as an osmoticum in osmotic adjustment experiments in these root tissues. Mannitol flows appear to be passive. Permeability to mannitol (about 10⁻¹⁰ m s⁻¹ is greater at the plasmalemma than at the tonoplast in carrot, and this would tend to cause the cytoplasm to swell. The implications for the control of cytoplasmic volume are discussed.     

Cadmium leads to stimulated expression of the lipid transfer protein genes in barley: implications for the involvement of lipid transfer proteins in wax assembly

In order to investigate the nature of genes expressed in leaf epidermal cells of higher plants, we have identified the nucleotide sequence of a cDNA designated ltp 7a2b encoding a novel nonspecific lipid transfer protein of barley (Hordeum vulgare L. cv. Gerbel). The cDNA of 755 basepairs contains an open reading frame of 366 nucleotides coding for a 12.3-kDa polypeptide. The first 29 amino acids constitute the putative signal peptide, characteristic for targeting to the secretory pathway. Analysis of mRNA levels by Northern blotting indicated that ltp 7a2b is preferentially expressed in the leaf epidermis. Levels of mRNA decreased during ageing of leaf tissue. Expression of ltp 7a2b was stimulated by a factor of 2–3 when the seedlings were grown in the presence of cadmium (10–1600 μM). Concomitantly, the primary leaves of Cd-exposed seedlings contained elevated levels of abscisic acid and a thicker wax layer of the cuticle. At 100 μM Cd in the hydroponic medium, the wax cover was increased by 50%. The increase in abscisic acid content, ltp 7a2b mRNA and wax coverage was either not seen, or seen much less, in Ni- and Zn-stressed seedlings. The data add circumstantial evidence to the recently proposed hypothesis that nonspecific lipid transfer proteins function in transfer of cutin and/or wax monomers from the site of synthesis in the cell to the cuticle. Received: 1 April 1996 / Accepted: 11 February 1997     

Regions associated with repression of the barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis>) <Emphasis Type="Italic">VERNALIZATION1</Emphasis> gene are not required for cold induction

Activity of the VERNALIZATION1 (VRN1) gene is required for flowering in temperate cereals such as wheat and barley. In varieties that require prolonged exposure to cold to flower (vernalization), VRN1 is expressed at low levels and is induced by vernalization to trigger flowering. In other varieties, deletions or insertions in the first intron of the VRN1 gene are associated with increased VRN1 expression in the absence of cold treatment, reducing or eliminating the requirement for vernalization. To characterize natural variation in VRN1, the first intron of the barley (Hordeum vulgare) VRN1 gene (HvVRN1) was assayed for deletions or insertions in a collection of 1,000 barleys from diverse geographical regions. Ten alleles of HvVRN1 containing deletions or insertions in the first intron were identified, including three alleles that have not been described previously. Different HvVRN1 alleles were associated with differing levels of HvVRN1 expression in non-vernalized plants and with different flowering behaviour. Using overlapping deletions, we delineated regions in the HvVRN1 first intron that are associated with low levels of HvVRN1 expression in non-vernalized plants. Deletion of these intronic regions does not prevent induction of HvVRN1 by cold or the maintenance of increased HvVRN1 expression following cold treatment. We suggest that regions within the first intron of HvVRN1 are required to maintain low levels of HvVRN1 expression prior to winter but act independently of the regulatory mechanisms that mediate induction of HvVRN1 by cold during winter. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nucleotide sequence data reported are available in the DDBJ/EMBL/GenBank databases under the accession numbers 1179825, 1179833, 1179836, 1179858.     

Supply of nitrogen can reverse senescence processes and affect expression of genes coding for plastidic glutamine synthetase and lysine-ketoglutarate reductase/saccharopine dehydrogenase

Nitrogen availability has a strong influence on developmental processes in plants. We show that the time of nitrogen supply regulates the course of leaf senescence in flag leaves of Hordeum vulgare . The senescence-specific decrease in chlorophyll content and photosystem II efficiency is clearly delayed when plants are fertilised with nitrate at the onset of leaf senescence. Concurrently, the additional supply of nitrate affects expression patterns of two marker genes of nitrogen metabolism. As shown by quantitative RT-PCR analyses, senescence-specific downregulation of plastidic glutamine synthetase ( GS2 ) and senescence-specific upregulation of lysine-ketoglutarate reductase/saccharopine dehydrogenase ( LKR/SDH ) are both clearly retarded. Depletion of nitrogen in experiments using hydroponic growth systems results in premature primary leaf senescence. The already started senescence processes can be even reversed by later nitrogen addition, as proved by a further increase in photosystem II efficiency and chlorophyll content, returning to the high values of controls which had not been deprived of nitrogen. Although both addition of nitrate or ammonium effectively reversed nitrogen depletion-induced primary leaf senescence, addition of urea did not. Additionally, effects of nitrogen supply on the course of leaf senescence were analysed in the model plant Arabidopsis thaliana. Leaves of A. thaliana show the same reversion of senescence processes after receiving additional nitrogen supply, indicating that the nitrogen response of leaf development is conserved in different plant species.     

Protease-stable integration of Lhcb1 into thylakoid membranes is dependent on chlorophyll b in allelic chlorina-f 2 mutants of barley (Hordeum vulgare L.)

Allelic chlorina-f2 mutants of barley (Hordeum vulgare L.) growing under different light and temperature conditions demonstrated that the chlorophyll b-free chlorina-f 2^{f  2} and chlorina-f 2¹⁰¹ express a stable phenotype. Only 3 out of 10 light-harvesting chlorophyll a/b-binding proteins, Lhca4 (photosystem I), and Lhcb1 and Lhcb6 (photosystem II), required chlorophyll b for accumulation. The other light-harvesting proteins were found in all chlorina-f2 alleles, indicating that the integration pathway of these proteins into mutant thylakoid membranes was not affected. Chlorina-f2 alleles with a thylakoid membrane capable of fullfilling photosynthesis and transport demands, but with various amounts of chlorophyll b: chlorina-f 2¹⁰¹ (chlorophyll b-free), chlorina-f2¹²³ (27% of chlorophyll b compared with the wild type) and chlorina-f 2¹²² (70% chlorophyll b), were chosen to investigate whether chlorophyll b is necessary for the protease-stable insertion of Lhcb l into mutant thylakoid membranes. The Lhcb1 was affected in almost all alleles and was most sensitive to chlorophyll b deficiency. The Lhcb1 antibody confirmed the heterogeneity of the polypeptides of the light-harvesting chlorophyll a/b-binding protein II (LHCII) and detected in wild-type membranes, two protease-resistant, mature forms of Lhcb1 with apparent molecular masses of 28 and 29 kDa. Only one band reacting with the Lhcb1 antibody could be detected in chlorophyll b-free chlorina-f 2  ^{f  2}. It co-migrated with the 29-kDa band, but was completely digested after treatment of the isolated mutant membranes with exogenous protease. This showed that in chlorina-f 2  f  ² the Lhcb1 precursor was processed at one cleavage site only. The resulting 29-kDa Lhcb1 was not provided with chlorophyll b, and, consequently, not properly folded and inserted into the membrane. It remained susceptible to protease and was inconvertable to a 28-kDa form. Received: 23 March 1998 / Accepted: 9 September 1998