Photosynthetic performance in wild emmer wheat,Triticum dicoccoides: ecological and genetic predictability

Summary In a twin study, we have shown that wild emmer wheat, Triticum dicoccoides, the progenitor of all cultivated wheats, harbours important genetic variation (Vg) in photosynthetic characteristics. This Vg resides within and between populations and ecogeographical regions in Israel, which is the center of origin and diversity of wild emmer wheat. Here we analyzed, by univariate and multivariate methods, the significant differentiation of variation in photosynthetic characteristics of 107 genotypes from 27 populations of wild emmer in Israel, distributed in three ecogeographical regions including central, xeric (northern cold and eastern warm) marginal, and mesic (western) marginal populations. The highest photosynthetic efficiency was displayed by populations of the xeric marginal region, but most variation for photosynthetic capacity occurs between accessions within ecogeographical regions and populations. Genotypes and populations of T. dicoccoides having high photosynthetic capacity can be identified by climatic factors and isozyme markers. The identification by genetic markers, if substantiated by testcrosses, can facilitate the maximization of conservation, in situ or ex situ, and utilization of these photosynthetic genetic resources for improvement of hexaploid wheat (T. aestivum).     

Genetic diversity for drought resistance in wild emmer wheat and its ecogeographical associations

Wild emmer wheat (Triticum turgidum spp. dicoccoides (Körn.) Thell.), the tetraploid progenitor of cultivated wheat, is a potential source for various agronomical traits, including drought resistance. The objectives of this study were to characterize (1) the genetic diversity for drought resistance in wild emmer wheat, and (2) the relationship between drought responses of the wild emmer germplasm and the ecogeographical parameters of its collection sites. A total of 110 wild emmer accessions consisting of 25 populations and three control durum wheat cultivars were examined under two irrigation regimes, well-watered (’wet’) and water-limited (’dry’). Wide genetic diversity was found both between and within the wild emmer populations in most variables under each treatment. A considerable number of the wild emmer accessions exhibited an advantage in productivity (spike and total dry matter) over their cultivated counterparts. Most wild emmer wheat accessions exhibited a greater carbon isotope ratio (δ¹³C, indicating higher water-use efficiency) under the dry treatment and higher plasticity of δ¹³C relative to the cultivated controls, which may have contributed to the drought adaptations in the former. The most outstanding drought-tolerance capacity (in term of productivity under the dry treatment and susceptibility indices) was detected in wild emmer populations originated from hot dry locations. The results suggest that wild emmer has the potential to improve drought resistance in cultivated wheat.     

Leaf characteristics and gas exchange of the mangrove <Emphasis Type="Italic">Laguncularia racemosa</Emphasis> as affected by salinity

Under constant salinity we analysed the leaf characteristics of Laguncularia racemosa (L.) Gaertn. in combination with gas exchange and carbon isotopic composition to estimate leaf water-use efficiency (WUE) and potential nitrogen-use efficiency (NUE). NaCl was not added to the control plants and the others were maintained at salinities of 15 and 30 ‰ (S₀, S₁₅, and S₃₀, respectively). Leaf succulence, sodium (Na), nitrogen (N), and chlorophyll (Chl) contents increased under salinity. Salinity had a negative impact on net photosynthetic rate (P _N) and stomatal conductance (g _s) at high and moderated irradiances. Potential NUE declined significantly (p<0.05) with salinity by 37 and 58 % at S₁₅ and S₃₀, respectively, compared to S₀ plants. Conversely, compared to S₀ plants, P _N/g _s increased under saline conditions by 12 % (S₁₅) and 50 % (S₃₀). Thus, WUE inferred from P _N/g _s was consistent with salinity improved short-term WUE. Long-term leaf WUE was also enhanced by salinity as suggested by significantly increased leaf δ¹³C with salinity. Improved WUE under salinity explains the eco-physiological success of mangrove species under increasing salinity. Conversely, decline in NUE may pose a problem for L. racemosa under hyper-saline environments regardless of N availability.     

Comparative ecophysiology of five species of Sedum (Crassulaceae) under well-watered and drought-stressed conditions

Summary Gas exchange patterns, diurnal malic acid fluctuations, and stable carbon isotope ratios of five species of Sedum were investigated to assess the ecophysiological characteristics of three different photosynthetic pathways under well-watered and drought-stressed conditions. All five species have succulent leaves and stems and were examined under identical environmental conditions. When well-watered, Sedum integrifolium (Raf.) Nels. and S. ternatum Michx. displayed C₃ photosynthesis, S. telephioides Michx. and S. nuttallianum Raf. exhibited CAM-cycling, and S. wrightii A. Gray showed CAM. When grown under a less frequent watering regime, S. integrifolium and S. ternatum exhibited CAM-cycling, whereas S. telephioides and S. nuttallianum displayed CAM-cycling simultaneously with low-level CAM. Sedum wrightii retained its CAM mode of photosynthesis. In general, leaf ¹³C values reflected these variations in photosynthetic pathways. While all values of water-use efficiency (WUE) were greater than those reported for most C₃ and C₄ species, no correlation of malic acid accumulation in the CAM and CAM-cycling (including low-level CAM) species with increased WUE was found. Sedum wrightii (CAM) had the highest WUE value at night, yet its 24-h WUE was not different from S. ternatum when the latter was in the C₃ mode. Thus, relative water-use efficiencies of these species of Sedum were not predictable based on photosynthetic pathways alone.     

Shade mitigates photoinhibition and enhances water use efficiency in kiwifruit under drought

We tested the hypotheses that a reduction of incident light of 50 % over sun-acclimated leaves of water stressed kiwifruit (Actinidia deliciosa var. deliciosa) would (1) reduce stomatal limitations to carbon supply and (2) mitigate the inactivation of the primary photochemistry associated with photosystem (PS) II, thereby this increases carbon gain and water-use efficiency (WUE). Groups of field-grown vines were either shaded or left naturally exposed and subjected to progressive water stress in order to study moderately and severely droughted vines, while other groups were well irrigated. Daily variations in leaf gas exchange and midday chlorophyll (Chl) a fluorescence were determined once plants had −0.6 MPa (moderate stress) and −1.0 MPa pre-down leaf water potential (severe stress). Variations in Chl pigment content and specific leaf area (SLA) are also discussed. Results reveal that 50 % shade application maintained efficiency of PSII close to 0.8 even under severe drought so that to prevent its large decline (0.65) recorded in sunlit leaves. Under moderate stress level stomata behaviour dominated upon metabolic impairments of PSII. Reduction of irradiance increased WUE (15–20 %) in droughted vines, representing a valuable intervention to save photosynthetic apparatus and improve WUE in vines experiencing typical Mediterranean summer stresses.     

Identification and characterization of a novel powdery mildew resistance gene <Emphasis Type="Italic">PmG3M</Emphasis> derived from wild emmer wheat, <Emphasis Type="Italic">Triticum dicoccoides</Emphasis>

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt) is one of the most important wheat diseases worldwide. Wild emmer wheat, Triticum turgidum ssp. dicoccoides, the tetraploid ancestor (AABB) of domesticated bread and durum wheat, harbors many important alleles for resistance to various diseases, including powdery mildew. In the current study, two tetraploid wheat mapping populations, derived from a cross between durum wheat (cv. Langdon) and wild emmer wheat (accession G-305-3M), were used to identify and map a novel powdery mildew resistance gene. Wild emmer accession G-305-3M was resistant to all 47 Bgt isolates tested, from Israel and Switzerland. Segregation ratios of F₂ progenies and F₆ recombinant inbred line (RIL) mapping populations, in their reactions to inoculation with Bgt, revealed a Mendelian pattern (3:1 and 1:1, respectively), indicating the role of a single dominant gene derived from T. dicoccoides accession G-305-3M. This gene, temporarily designated PmG3M, was mapped on chromosome 6BL and physically assigned to chromosome deletion bin 6BL-0.70-1.00. The F₂ mapping population was used to construct a genetic map of the PmG3M gene region consisted of six simple sequence repeats (SSR), 11 resistance gene analog (RGA), and two target region amplification polymorphism (TRAP) markers. A second map, constructed based on the F₆ RIL population, using a set of skeleton SSR markers, confirmed the order of loci and distances obtained for the F₂ population. The discovery and mapping of this novel powdery mildew resistance gene emphasize the importance of the wild emmer wheat gene pool as a source for crop improvement.     

Genetic diversity of wild emmer wheat in Israel and Turkey

Summary Allozyme variation in the tetraploid wild emmer wheat, Triticum dicoccoides, the progenitor of all cultivated wheats, was studied for the proteins encoded by 42 gene loci in 1815 plants representing 37 populations - 33 from Israel and 4 from Turkey - sampled in 33 localities from 1979 to 1987. The results showed that: (a) 6 loci (14%) were monomorphic in all populations, 15 loci (36%) were locally polymorphic, and 21 loci (50%) were regionally polymorphic. These results are similar to those obtained earlier on 12 Israeli populations. All polymorphic loci (except 4) displayed high local levels of polymorphism (>/ 10%). (b) The mean number of alleles per locus, A, was 1.252 (range: 1.050–1.634); the proportion of polymorphic loci per population averaged 0.220 (range: 0.050–0.415); genic diversity, He, averaged 0.059 (range: 0.002–0.119). (c) Altogether there were 119 alleles at the 42 putative loci tested, 114 of these in Israel, (d) Genetic differentiation was primarily regional and local, not clinal; 70% of the variant alleles were common (>/ 10%) and not widespread, but rather localized or sporadic, displaying an archipelago population genetics and ecology structure. The coefficients of genetic distance between populations were high and averaged D = 0.134; range: 0.018–0.297, an indication of sharp genetic differentiation over short distances, (e) Discriminant analyses differentiated Israeli from Turkish populations, and within Israel, between central and 3 marginal regions, as well as between different soil-type populations, (f) Allozymic variation comprised 40% within and 60% between populations, (g) Gametic phase disequilibria were abundant, their number being positively correlated (r_s = 0.60, P<0.01) with the humidity, (h) Multilocus organization was substantive, also positively correlated with humidity, (i) Allozyme diversity, overall and at single loci, was significantly correlated with, and partly predictable by, climatic and edaphic factors, (j) The distrubition of the significant positive and negative values and the absence of autocorrelations in the correlogram revealed no similar geographic patterns across loci, eliminating migration as a prime factor of population genetic differentiation. These results suggest: (I) during the evolutionary history of wild emmer, diversifying natural selection, through climatic and edaphic factors, was a major agent of genetic structure and differentiation at both the single and multilocus levels; (II) wild emmer harbors large amounts of genetic diversity exploitable as genetic markers in sampling and abundant genetic resources utilizable for wheat improvement.     

Changes of the photosynthetic behaviour in annual C<Subscript>3</Subscript> species at late successional stage under environmental drought conditions

Differences in structural, physiological, and biochemical features between C₃ and C₄ species resulted in different wateruse efficiencies and different adaptations to climate. This paper aimed at investigating, at a late successional stage, the water-use efficiency of two forage species, Dichanthium ischaemum and Dasypyrum villosum, which exhibit different growth forms (perenial, annual) and photosynthetic mechanisms (C₄ and C₃, respectively). The annual C₃ species Avena fatua, at an early successional stage, was included in our experiments to contrast its behaviour against D. villosum. The experiment was conducted during the growing season in low-elevation grasslands of North Greece. Midday leaf water potential, net photosynthetic rate, transpiration rate and stomatal conductance were measured. Instantaneous water-use efficiency (WUE) and intrinsic water-use efficiency (WUEi) were calculated in D. ischaemum, D. villosum, and A. fatua. The results suggest that, under natural rainfall conditions, the annual C₃ grass species D. villosum exhibits a similar WUE with higher values of WUE_i than the perennial C₄ species D. ischaemum at late stage of succession on the low elevation Mediterranean grasslands. Moreover, A. fatua at an early successional stage, exhibited different photosynthetic behaviour than D. villosum at a late successional stage. These findings indicate that the annual C₃ species D. villosum under drought and at a late successional stage seems to modify the WUE obtaining values similar to those of C₄ species. The extent to which the ecophysiological characteristics of D. villosum are environmentally or intrinsically determined remains to be answered.     

Variations of leaf morphology,photosynthetic traits and water-use efficiency in Western-Mediterranean tomato landraces

Modern tomato (Solanum lycopersicum L.) breeding has mainly focused on increasing productivity under unlimited watering. In contrast, some Mediterranean accessions have been traditionally cultivated under water shortage and selected on the basis of their water-use efficiency (WUE). Ramellet and Penjar landraces were planted with other traditional, old and modern inbreeds, under full irrigation. In order to found differences between the tomato accessions, gas-exchange and leaf morphology measurements were performed. Despite high variability, Ramellet and Penjar presented clear differences compared to modern cultivars, mostly related to leaf morphology and photosynthetic traits, while no differences were found in WUE. Results highlighted that better leaf CO₂ conductance might be a main factor determining the improvement of net CO₂ assimilation and WUE.     

Photosynthesis,water-use efficiency and δ<Superscript>13</Superscript>C of five cowpea genotypes grown in mixed culture and at different densities with sorghum

A field experiment involving two planting densities (83,333 and 166,666 plants per ha), two cropping systems (monoculture and mixed culture) and five cowpea [Vigna unguiculata L. (Walp.)] genotypes was conducted at Nietvoorbij (33°54S, 18°14E), Stellenbosch, South Africa, to select cowpea material with superior growth and water-use efficiency (WUE). The results showed significantly higher photosynthetic rates, stomatal conductance and transpiration in leaves of plants at low density and in monoculture due to greater chlorophyll (Chl) levels relative to those at high density and in mixed culture. As a result, C concentration in leaves and the amount of C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B accumulated in shoots at low density and under monoculture were also much higher. Even though no marked differences in photosynthetic rates were found between and among the five cowpea genotypes, leaf C concentration and shoot C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B contents differed considerably, with Sanzie exhibiting the highest C concentration and C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B contents in shoots, followed by Bensogla and Omondaw, while ITH98-46 and TVu1509 had the lowest shoot concentration and contents of C, P, K, Ca, Mg, Fe, Cu, Zn, Mn, and B. WUE (calculated as photosynthate produced per unit water molecule transpired) was significantly greater in plants at low density and monoculture relative to those at high density and in mixed culture. Isotopic analysis revealed significant differences in δ¹³C values of sorghum [Sorghum bicolor L. (Moench.)] and cowpea, with higher δ¹³C values being obtained for plants at low density and in monoculture relative to those at high density or in mixed culture. The five cowpea genotypes also showed significant differences in δ¹³C values, with Sanzie exhibiting the most negative value (i.e. low WUE) and ITH98-46, the least negative δ¹³C value (i.e. high WUE). Whether measured isotopically or from gas-exchange studies, sorghum (a C₄ species) exhibited much higher WUE relative to cowpea (a C₃ species). Both correlation and regression analyses revealed a positive relationship between WUE from gas-exchange studies and δ¹³C values from isotopic analysis of cowpea and sorghum shoots.     

Variance for water-use efficiency among ecotypes and recombinant inbred lines of Arabidopsis thaliana (Brassicaceae)

Knowledge of natural variation among ecotypes and recombinant inbred lines of Arabidopsis thaliana L. Heynh for season-long water-use efficiency (WUE, moles of carbon accumulated per mole of water used) is useful in the design of experiments to understand the genetic control of this phenomenon. Water-use efficiency among 31 container-grown Arabidopsis ecotypes ranged from a high of 2.40 to a low of 1.86 mg/g (grams of dry aerial biomass per gram of water used). Genetic variance for WUE was observed among 80 F₅ recombinant inbred lines (RILs) derived from a cross between the highest (Lip-0) by the lowest (Edi-0) ecotype. The heritability of WUE was calculated as 0.18 ± 0.07. The mean WUE for the ten highest and ten lowest RILs was 2.42 and 1.05 mg/g, respectively. A significant difference was observed in the composition of stable isotopes of plant carbon (δ¹³C) between the mean of the ten highest (-31.23±) and the mean of the ten lowest (-31.96±) RILs based on WUE. Characterization of the 80 RILs provides a structured population for further genetic and physiologic study of WUE.     

Genetic analysis of photosynthetic variation in hexaploid and tetraploid wheat and their interspecific hybrids

Intra- and inter-specific variation in CO₂ assimilation rate (A) in Triticum spp. is well documented for reproductive growth stages. Research was conducted to characterize early vegetative photosynthetic variation in a diverse set of cultivated hexaploid wheat (T. aestivum L.) germplasm and in wild tetraploid (T. dicoccoides Korn) and hexaploid x tetraploid populations. Choice of hexaploid genotypes was based on maximum genetic distance between cultivars within the HRW and SRW wheat classes of the USA. The tetraploid material was produced by hybridizing two accessions of T. dicoccoides previously shown to differ widely in A and A/Chl but with similar leaf morphology. Genetic variability in the HRW and SRW gene pools was attributed to more recently developed descendent lines and unrelated lines rather than parental lines. Phenotypic distributions for A, stomatal conductance (gs), and internal CO₂ concentration (Ci) in the F₂ tetraploid population were continuous and showed transgressive segregation, reflecting quantitative inheritance with intermediate heritability. Variability in A was not associated with chlorophyll content or CO₂ supply to the mesophyll measured as Ci. Genetic variability in A was also observed in the interspecific backcross population, 2*TAM W-101/PI 428109, thereby providing a germplasm pool to select for high A while restoring the D genome of hexaploid wheat. These results suggest that genetic improvement of vegetative assimilation rate is feasible in hexaploid wheat via homologous transfer from an alien source.Abbreviations HRW hard red winter - LA leaf area - r_G genotypic correlation - r_P phenotypic correlation - SRW soft red winter     

Growth, gas exchange, water-use efficiency, and carbon isotope composition of ‘Gale Gala’ apple trees grafted onto 9 wild Chinese rootstocks in response to drought stress

To determine the effects of rootstock choice on the scion response to drought stress, we compared the vegetative growth, biomass accumulation, gas exchange, and water-use efficiency (WUE) of ??Gale Gala?? apple (Malus domestica Borkh.) trees grafted onto nine wild Chinese Malus rootstocks. Compared with the well-watered control, drought treatment limited growth, as manifested by smaller increments in plant height (PH), trunk diameter (TD), total fresh biomass (TB), total dry biomass (TDB), total leaf area (LA), and relative growth rate (RGR). The extent of this effect differed among rootstocks. Stress conditions led to increases in the root/shoot ratio (RSR), leaf thickness (LT), water-holding capacity (WHC), carbon isotope composition (??¹³C), and WUE. Decreases were noted in stomatal density (SD), leaf relative water content (RWC), chlorophyll content (Chl), net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s), again varying by rootstock. Those that are generally considered more drought-tolerant, e.g., M. sieversii, M. prunifolia, and M. toringoides, had smaller declines in PH, TD, TB, TDB, LA, RGR, SD, RWC, Chl, P _N, E, and g _s and proportionally greater increases in RSR, LT, WHC, ??¹³C, and WUE compared with the droughtsensitive M. hupehensis and M. sieboldii. These results suggest that moisture stress has a significant dwarfing effect in the latter two species. Based on WUE calculations, trees on drought-tolerant rootstocks showed higher tolerance when stressed, whereas those on drought-sensitive rootstocks were less tolerant, as indicated by their lower WUE values.     

Differential responses of leaf water-use efficiency and photosynthetic nitrogen-use efficiency to fertilization in Bt-introduced and conventional rice lines

Leaf stomatal conductance (g _s), transpiration rate (E), and light-saturated net photosynthetic rate (P _Nmax) at three developmental stages (tillering stage, jointing-booting stage, and milking stage) and leaf total nitrogen concentration (LTNC) and δ¹³C value at milking stage were measured for a conventional rice line (Minghui 63) and its corresponding Bacillus thuringiensis (Bt)-gene [cry1A (b and c)] introduced line (Bt line) under three fertilizer levels. Compared to conventional line, Bt line showed lower g _s, which was associated with lower P _Nmax and E, but instantaneous water-use efficiency (WUE), measured as the ratio of P _Nmax to E, was higher in the Bt line than in the conventional line, particularly in the jointing-booting stage. However, δ¹³C values were not significantly different across treatments, suggesting that intrinsic water-use efficiency (WUE_in) might be indistinguishable between Bt and conventional lines. LTNC was higher but P _Nmax was lower in Bt line compared to conventional line, resulting in significantly lower photosynthetic nitrogen-use efficiency (PNUE). This might result from the additional cost of producing Bt protein in the Bt line due to the effect of competing nitrogen with photosynthetic machinery. Bt-gene introduction and expression does not significantly change WUE_in but may significantly decrease leaf PNUE. Thus we suggest that Bt rice should be carefully examined in relation to environmental risks (e.g. water-body pollution) before planting commercially.     

Water-stress and N-nutrition effects on photosynthesis and growth of Brassica carinata

Effects of water and nitrogen (N) supply on growth and photosynthetic response of B. carinata were examined in this study. Plant growth and related characteristics varied significantly in response to the availability of water and nitrogen. B. carinata maximized the utilization efficiency of the most limiting resources by developing physiological adaptations, such as changes in root and leaf development. The utilization of water and N was tightly linked with the availability of each resource. Instantaneous water-use efficiency (WUE) was always greater in plants with high-N nutrition [50, 100, and 150 kg(N) ha^?1] than in the low-N-treated plants [0 kg(N) ha^?1] in all watering treatments. Instantaneous N-use efficiency (PNUE) decreased significantly with increasing water stress in all N treatments. Seed yield is significantly related to PNUE (p>0.05) but not WUE (p<0.05). The positive relationship between leaf net photosynthetic rate (P _N) and seed yield suggests that P _N can be used as an important tool for selection of new strains with high seed yield.     

Genetic variation in and covariation between leaf gas exchange,morphology, and development in Polygonum arenastrum,an annual plant

Summary We present evidence of genetic variation in and covariation between leaf-level gas exchange properties and leaf size among family lines of Polygonum arenastrum. This self-fertilizing annual had previously been shown to vary genetically in developmental phenology and in morphology (size of leaves, internodes, flowers and seeds) (Geber 1990). Significant family differences were found in photosynthetic carbon assimilation rate (A), lcaf conductance to water vapor (g), instantaneous water-use efficiency (WUE), and leaf carbon isotope discrimination (). A strong positive genetic correlation between A and g suggested that there was stomatal limitation on A. In addition, higher g led to relatively greater increases in transpiration, E, than in assimilation, A, so that families with high rates of gas exchange had lower instantaneous WUE and/or higher carbon isotope discrimination values. Leaf size and gas exchange were genetically correlated. In earlier studies leaf size was found to be genetically correlated with developmental phenology (Geber 1990). The pattern that emerges is one in which small-leaved families (which also have small internodes, flowers, and seeds) tend to have high gas exchange rates, low WUE, rapid development to flowering and high early fecundity, but reduced life span and maximum (vegetative and reproductive) yield compared to large-leaved families. We suggest that this pattern may have arisen from selection for contrasting suites of characters adapted to environments differing in season length.     

Responses of photosynthesis and carbohydrate-partitioning to limitations in nitrogen and water availability in field-grown sunflower*

Abstract. Sunflower plants (Helianthus annuus L., cv. CGL 208) were field-grown in adjacent plots of varying resource availability. Control plants received irrigation (on a 4–5 d interval) and high levels of fertilizer nitrogen. Nutrient-stress (N-stress) plants received control levels of irrigation but no nutrient amendments and were determined to be nitrogen-limited. Water-stress (H₂O-stress) plants received control levels of fertilizer nitrogen, but no irrigation after approximately 6 weeks of plant growth. Both stress treatments reduced maximum and diurnal net photosynthesis (A) but resulted in different physiological or biochemical adjustments that tended to maintain or increase A per unit of resource (nitrogen or water) in shortest supply while decreasing the ratio of A per unit of abundant resource. Nutrient-stress reduced total foliar nitrogen, foliar chlorophyll, and initial and total RuBPCase activities, thereby enhancing or preserving photosynthetic nitrogen-use efficiency (NUE), defined as the maximum A observed per unit of leaf nitrogen, relative to the control and H₂O-stress treatments. In addition, N-stress reduced photosynthetic water-use efficiency (WUE), defined as the ratio of A to stomatal conductance to water vapour (g). The slope of A versus g increased with H₂O-stress. In addition, sunflower plants responded to H₂O-stress by accumulating foliar glucose and sucrose and by exhibiting diurnal leaf wilting, which presumably provided additional improvements in photosynthetic WUE through osmoregulation and reduction of midday radiation interception respectively. Photosynthetic NUE was decreased by H₂O-stress in that control levels of total nitrogen, foliar chlorophyll, and RuBPCase activities were maintained even after mean diurnal levels of A had fallen to less than 50% of the control level. We conclude that field-grown sunflower manages a trade-off between photosynthetic WUE and NUE, increasing use efficiency of the scarce resource while decreasing use efficiency of the abundant resource.     

Geographical variation in heading traits in wild emmer wheat, Triticum dicoccoides. I. Variation in vernalization response and ecological differentiation

 Geographical variation in vernalization response and narrow-sense earliness was investigated for accessions of wild emmer wheat, Triticum dicoccoides, collected in Israel. Wide variation between and within populations was observed in both characters. The analysis of vernalization response showed that 2 accessions from Tabigha were of a strong spring growth habit, and thus wild emmer wheat was classified into four types, i.e., strongly spring type, moderately spring type, moderately winter type, and strongly winter type, according to their vernalization response. Whereas winter types were frequently found in most populations except that of Tabigha, the distribution of spring types was sporadic and restricted to warmer areas. It was thus suggested that spring type in T. dicoccoides might have evolved from a winter prototype as an adaptation to warmer conditions. Within moderately winter and moderately spring types, quantitative differences in vernalization response, measured as Dof₇₀/Dof₂₀ and Dof₂₀/Dof₀, were observed between populations. Inter- and intra-population variation in vernalization response could be explained to some extent by the difference in growing conditions at each habitat. It was clearly indicated that environmental heterogeneity caused ecogenetic differentiation in wild emmer wheat in Israel. Wild emmer wheat also varied considerably for narrow-sense earliness, ranging from 32.9 days to 69.5 days among accessions. However, it was difficult to explain its geographical variation simply by a linear relationship with environmental factors, and a nonlinear relationship and/or unknown microgeographic heterogeneity may be responsible. Received: 18 March 1996/Accepted: 13 December 1996     

Effects of fertilization on leaf photosynthetic characteristics and grain yield in tartary buckwheat Yunqiao1

Tartary buckwheat (Fagopyrum tataricum Gaertn) has been praised as one of green foods for humans in the 21_st century. Effects of fertilization on leaf photosynthetic characteristics and grain yield of tartary buckwheat has not been yet reported in detail. Our experiment was set as a split-plot factorial. The main plots and subplots were designed by fertilizer ratio and rate as: NPK 1:1:1 (A1), NPK 1:4:2 (A2), NPK 1:2:3 (A3), and 300 (B1), 450 (B2), and 600 (B3) kg (NPK) ha^–1. Our results showed that the grain yield was significantly and positively correlated with the net photosynthetic rate (P _N), stomatal conductance (g _s), transpiration rate (E), PAR, stomatal limitation value (L_s), chlorophyll content (SPAD value), and leaf area index (LAI), while significantly and negatively correlated with intercellular CO₂ concentration (C _i) and water-use efficiency (WUE). The grain yield, P _N, g _s, E, PAR, L_s, SPAD, and LAI increased and then decreased with enhanced fertilization, and their maximum values appeared in the A2B2 treatment. The C _i and WUE decreased and then increased with enhanced fertilization, and their minimum values appeared in the A2B2 treatment. Our results suggested that fertilization had significant effects on the leaf photosynthetic capacity and grain yield of tartary buckwheat Yunqiao 1, and the best fertilization strategy was 450 kg ha^–1 with NPK 1:4:2.