Plant factors controlling seed set in maize : the influence of silk, pollen, and ear-leaf water status and tassel heat treatment at pollination

In maize (Zea mays L.) large decreases in kernel number result when low water potentials (Ψ_w) and high temperatures occur during pollination. To gain insight into the basis for the decreased seed set, silk, pollen, and ear-leaf Ψ_w, the capability for silk osmotic adjustment, and pollen appearance were measured to determine their relationship to seed set. A multiple-eared or prolific (high carbohydrate availability to the pistillate inflorescence) hybrid (B73 × FR25), a heat sensitive hybrid (WF9 × A632), and a commercial hybrid (B73 × Mo17) were studied. A cross-pollination experiment, with pollination limited by pollen amount, was conducted to determine the impact on seed set of water and heat stressing the tassel and water stressing the ear. At low Ψ_w, silk Ψ_w and seed set were decreased whereas pollen Ψ_w, appearance, and viability were unaffected. High temperature resulted in a 2 megapascal decrease in pollen Ψ_w, visually damaged pollen being shed, decreased pollen viability, and, in two of the hybrids, substantially decreased pollen shed. Prolificacy did not result in increased silk solute accumulation but did result in superior seed production by the pistillate inflorescence at low Ψ_w. The magnitude of the decrease in silk solute potential was small (0.2 megapascal) and similar for all genotypes. One hybrid maintained a relatively high silk turgor but this hybrid also decreased the most in seed production when the pistillate inflorescence was water deficient. These results indicated an adverse effect of high temperature on pollen development, a positive relationship between seed production and silk water status, and no advantage to high silk turgor after silk emergence in maintaining seed production. Additionally, there was no evidence of variation in silk solute regulation capability among hybrids which varied in prolificacy, a trait important in drought tolerance, but the seed production of the pistillate inflorescence of the prolific hybrid was least affected by water deficit.     

Correlation between the Maintenance of Photosynthesis and in Situ Protoplast Volume at Low Water Potentials in Droughted Wheat

Studies were undertaken to examine the relationship between water deficit effects on photosynthesis and the extent of protoplast volume reduction which occurs in leaves at low water potential (Ψ_w). This relationship was monitored in two cultivars (`Condor' and `Capelle Desprez') of cultivated wheat (Triticum aestivum) that differed in sensitivity to drought, and in a wild relative of cultivated wheat (Triticum kotschyi) that has been previously found to be `drought resistant.' When subjected to periods of water stress, Condor and T. kotschyi plants underwent osmotic adjustment; Capelle plants did not. Photosynthetic capacity was maintained to different extents in the three genotypes as leaf Ψ_w declined during stress; Capelle plants were most severely affected. Calculations of internal leaf [CO₂] and stomatal conductance from gas exchange measurements indicated that differences in photosynthetic inhibition at low Ψ_w among the genotypes were primarily due to nonstomatal effects. The extent of protoplast volume reduction that occurred in leaves at low Ψ_w was also found to be different in the three genotypes; maintenance of protoplast volume and photosynthetic capacity in stressed plants of the genotypes appeared to be correlated. When the extent of water stress-induced inhibition of photosynthesis was plotted as a function of declining protoplast volume, this relationship appeared identical for the three genotypes. It was concluded that there is a correlative association between protoplast volume and photosynthetic capacity in leaves of wheat plants subjected to periods of water stress.     

Osmotic Adjustment in Sorghum: II. Relationship to Gas Exchange Rates

Lowering of the solute potential by osmotic adjustment (OA) has been proposed to allow maintenance of leaf turgor potential (Ψ_p), stomatal conductance (g), and photosynthesis (A) at low leaf water potential. However, literature concerning the role of OA in the maintenance of g and A under water stress is limited and often contradictory. The objective of this experiment was to examine the association of OA with g and A in grain sorghum (Sorghum bicolor L. Moench). A single sorghum hybrid (cv AT×623 × RT×430) was studied under field conditions using four different water supplies. Diurnal and midday water potential, solute potential, Ψ_p, OA, g, and A were measured during preflowering and grain-filling growth stages. A second experiment was conducted under greenhouse conditions. Two sorghum genotypes (BT×623 and BT×378) differing in their g and A responses to plant water stress were compared for their OA capacity during a water deficit cycle imposed from the beginning of panicle initiation through flowering. Under both field and greenhouse conditions, g and A rapidly declined with increased water stress despite the occurrence of OA. Under greenhouse conditions, BT×623 maintained significantly higher g and A than BT×378 during the water stress cycle. However, no significant differences in OA or Ψ_p existed between the two genotypes, indicating that OA was not associated with differences observed in g and A between these genotypes. We conclude that the response of g and A to water stress was not directly associated with OA and certainly was not maintained by OA.     

Carbon isotope discrimination varies genetically in c(4) species

Carbon-isotope discrimination (Δ) is used to distinguish between different photosynthetic pathways. It has also been shown that variation in Δ occurs among varieties of C₃ species, but not as yet, in C₄ species. We now report that Δ also varies among genotypes of sorghum (Sorghum bicolor Moench), a C₄ species. The discrimination in leaves of field-grown plants of 12 diverse genotypes of sorghum was measured and compared with their grain yields. Discrimination varied significantly among genotypes, and there was a significant negative correlation between grain yield and Δ. The variation in Δ may be caused by genetic differences in either leakiness of the bundle-sheath cells or by differences in the ratio of assimilation rate to stomatal conductance. At the leaf level, the former should be related to light-use efficiency of carbon fixation and the latter should be related to transpiration efficiency. Both could relate to the yield of the crop.     

Chloroplast osmotic adjustment and water stress effects on photosynthesis

Previous studies have suggested that chloroplast stromal volume reduction may mediate the inhibition of photosynthesis under water stress. In this study, the effects of spinach (Spinacia oleracea, var `Winter Bloomsdale') plant water deficits on chloroplast photosynthetic capacity, solute concentrations in chloroplasts, and chloroplast volume were studied. In situ (gas exchange) and in vitro measurements indicated that chloroplast photosynthetic capacity was maintained during initial leaf water potential (Ψ_w) and relative water content (RWC) decline. During the latter part of the stress period, photosynthesis dropped precipitously. Chloroplast stromal volume apparently remained constant during the initial period of decline in RWC, but as leaf Ψ_w reached −1.2 megapascals, stromal volume began to decline. The apparent maintenance of stromal volume over the initial RWC decline during a stress cycle suggested that chloroplasts are capable of osmotic adjustment in response to leaf water deficits. This hypothesis was confirmed by measuring chloroplast solute levels, which increased during stress. The results of these experiments suggest that stromal volume reduction in situ may be associated with loss of photosynthetic capacity and that one mechanism of photosynthetic acclimation to low Ψ_w may involve stromal volume maintenance.     

Leaf water relations and maintenance of gas exchange in coffee cultivars grown in drying soil

Plant water status, leaf tissue pressure-volume relationships, and photosynthetic gas exchange were monitored in five coffee (Coffea arabica L.) cultivars growing in drying soil in the field. There were large differences among cultivars in the rates at which leaf water potential (Ψ_L) and gas exchange activity declined when irrigation was discontinued. Pressure-volume curve analysis indicated that increased leaf water deficits in droughted plants led to reductions in bulk leaf elasticity, osmotic potential, and in the Ψ_L at which turgor loss occurred. Adjustments in Ψ_L at zero turgor were not sufficient to prevent loss or near loss of turgor in three of five cultivars at the lowest values of midday Ψ_L attained. Maintenance of protoplasmic volume was more pronounced than maintenance of turgor as soil drying progressed. Changes in assimilation and stomatal conductance were largely independent of changes in bulk leaf turgor, but were associated with changes in relative symplast volume. It is suggested that osmotic and elastic adjustment contributed to maintenance of gas exchange in droughted coffee leaves probably through their effects on symplast volume rather than turgor.     

Variation in climatic tolerance,but not stomatal traits,partially explains Pooideae grass species distributions

Background and AimsGrasses in subfamily Pooideae live in some of the world’s harshest terrestrial environments, from frigid boreal zones to the arid windswept steppe. It is hypothesized that the climate distribution of species within this group is driven by differences in climatic tolerance, and that tolerance can be partially explained by variation in stomatal traits.MethodsWe determined the aridity index (AI) and minimum temperature of the coldest month (MTCM) for 22 diverse Pooideae accessions and one outgroup, and used comparative methods to assess predicted relationships for climate traits versus fitness traits, stomatal diffusive conductance to water (g_w) and speed of stomatal closure following drought and/or cold.Key ResultsResults demonstrate that AI and MTCM predict variation in survival/regreening following drought/cold, and g_w under drought/cold is positively correlated with δ ¹³C-measured water use efficiency (WUE). However, the relationship between climate traits and fitness under drought/cold was not explained by g_w or speed of stomatal closure.ConclusionsThese findings suggest that Pooideae distributions are at least partly determined by tolerance to aridity and above-freezing cold, but that variation in tolerance is not uniformly explained by variation in stomatal traits.     

Osmotic adjustment, symplast volume, and nonstomatally mediated water stress inhibition of photosynthesis in wheat

At low water potential (ψ_w), dehydration reduces the symplast volume of leaf tissue. The effect of this reduction on photosynthetic capacity was investigated. The influence of osmotic adjustment on this relationship was also examined. To examine these relationships, comparative studies were undertaken on two wheat cultivars, one that osmotically adjusts in response to water deficits (`Condor'), and one that lacks this capacity (`Capelle Desprez'). During a 9-day stress cycle, when water was withheld from plants grown in a growth chamber, the relative water content of leaves declined by 30% in both cultivars. Leaf osmotic potential (ψ_s) declined to a greater degree in Condor plants. Measuring ψ_s at full turgor indicated that osmotic adjustment occurred in stressed Condor, but not in Capelle plants. Two methods were used to examine the degree of symplast (i.e. protoplast) volume reduction in tissue rapidly equilibrated to increasingly low ψ_w. Both techniques gave similar results. With well-watered plants, symplast volume reduction from the maximum (found at high ψ_w for each cultivar) was the same for Condor and Capelle. After a stress cycle, volume was maintained to a greater degree at low ψ_w in Condor leaf tissue than in Capelle. Nonstomatally controlled photosynthesis was inhibited to the same degree at low ψ_w in leaf tissue prepared from well-watered Condor and Capelle plants. However, photosynthetic capacity was maintained to a greater degree at low ψ_w in tissue prepared from stressed Condor plants than in tissue from stressed Capelle plants. Net CO₂ uptake in attached leaves was monitored using an infrared gas analyzer. These studies indicated that in water stressed plants, photosynthesis was 106.5% higher in Condor than Capelle at ambient [CO₂] and 21.8% higher at elevated external [CO₂]. The results presented in this report were interpreted as consistent with the hypothesis that there is a causal association between protoplast (and presumably chloroplast) volume reduction at low ψ_w and low ψ_w inhibition of photosynthesis. Also, the data indicate that osmotic adjustment allows for maintenance of relatively greater volume at low ψ_w, thus reducing low ψ_w inhibition of chloroplast photosynthetic potential.     

Transpiration- and growth-induced water potentials in maize

Recent evidence from leaves and stems indicates that gradients in water potential (ψ_w) necessary for water movement through growing tissues are larger than previously assumed. Because growth is sensitive to tissue ψ_w and the behavior of these gradients has not been investigated in transpiring plants, we examined the water status of all the growing and mature vegetative tissues of maize (Zea mays L.) during high and low rates of transpiration. The ψ_w measured in the mature regions of the plant responded primarily to transpiration, while the ψ_w in the growing regions was affected both by transpiration and growth. The transpiration-induced potentials of the mature tissue formed a gradient of decreasing ψ_w along the transpiration stream while the growth-induced potentials formed a gradient of decreasing ψ_w from the transpiration stream to the expanding cells in the growing tissue. The growth-induced gradient in ψ_w within the leaf remained fairly constant as the xylem ψ_w decreased during the day and was associated with a decreased osmotic potential (ψ_s) of the growing region (osmotic adjustment). The growth-induced gradient in ψ_w was not caused by excision of the tissue because intact maize stems exhibited a similar ψ_w. These observations support the concept that large gradients in ψ_w are required to maintain water flow to expanding cells within all the vegetative tissues and suggest that the maintenance of a favorable gradient in ψ_w for cell enlargement may be an important role for osmotic adjustment.     

Effect of water deficits on seed development in soybean : I. Tissue water status

Water deficits during seed filling often decrease seed size in soybean (Glycine max L.). The physiological basis for this response is not known but may result from direct effects of low seed water potential (Ψ_w) on the seed filling process. To determine whether low Ψ_w occurred in reproductive tissues of soybean, we monitored the water status (Ψ_w, Ψ_s, and Ψ_p) of leaf, pericarp, and seed (embryo and testa) tissue of greenhouse-grown plants subjected to a brief water deficit during the linear period of seed growth. Water deficits were imposed by withholding water and monitored in the reproductive tissues by thermocouple psychrometry. When water was abundant, leaf, pericarp, and seed Ψ_w were −0.5 to −0.7 megapascal at midday. When water was withheld, leaf Ψ_w decreased to −2.3 megapascals within 6 days. Pericarp Ψ_w also decreased to −1.9 megapascal during this time. Pericarp Ψ_s followed the decline in Ψ_w, but osmotic adjustment was not evident as the pericarp lost turgor completely by day 6. However, seed Ψ_w, Ψ_s, and Ψ_p were not significantly different from the controls. These results indicate that the water status of the developing seeds of soybean is not altered by short-term water deficits severe enough to inhibit the metabolic activity of the maternal plant. Maintenance of a favorable water status may be important for the conservation of seed growth rate exhibited by soybean under dry conditions.     

Genetic variation in stomatal and biochemical limitations to photosynthesis in the annual plant, Polygonum arenastrum

 Terrestrial plant photosynthesis may be limited both by stomatal behavior and leaf biochemical capacity. While inferences have been made about the importance of stomatal and biochemical limitations to photosynthesis in a variety of species in a range of environments, genetic variation in these limitations has never been documented in wild plant populations. Genetic variation provides the raw material for adaptive evolution in rates of carbon assimilation. We examined genetic variation in gas exchange physiology and in stomatal and biochemical traits in 16 genetic lines of the annual plant, Polygonum arenastrum. The photosynthesis against leaf internal CO₂ (A−ci) response curve was measured on three greenhouse-grown individuals per line. We measured the photosynthetic rate (A) and stomatal conductance (g), and calculated the internal CO₂ concentration (ci) at ambient CO₂ levels. In addition, the following stomatal and biochemical characteristics were obtained from the A−ci curve on each individual: the degree of stomatal limitation to photosynthesis (L_s), the maximum ribulose 1,5-biphosphate carboxylase-oxygenase (Rubisco) activity (Vc_max) and electron transport capacity (J_max). All physiological traits were genetically variable, with broad sense heritabilities ranging from 0.66 for L_s to 0.94 for J_max. Strong positive genetic correlations were found between Vc_max and J_max, and between g and biochemical capacity. Path analyses revealed strong causal influences of stomatal conductance and leaf biochemistry on A and ci. Path analysis also indicated that L_s confounds both stomatal and biochemical effects, and is an appropriate measure of stomatal influences on photosynthesis, only when biochemical variation is accounted for. In total, our results indicate that differences among lines in photosynthesis and ci result from simultaneous changes in biochemical and stomatal characteristics and are consistent with theoretical predictions that there should be co-limitation of photosynthesis by ribulose-1,5-biphosphate (RuBP) utilization and regeneration, and by stomatal conductance and leaf biochemistry. Gas exchange characteristics of genetic lines in the present study were generally consistent with measurements of the same lines in a previous field study. Our new results indicate that the mechanisms underlying variation in gas exchange include variation in both stomatal conductance and biochemical capacity. In addition, A, g, and ci in the present study tended also to be positively correlated with carbon isotope discrimination (Δ), and negatively correlated with time to flowering, life span, and leaf size based on earlier work. The pattern of correlation between physiology and life span among genetic lines of P. arenastrum parallels interspecific patterns of character correlations. We suggest that the range of trait constellations among lines in P. arenastrum represents a continuum between stress avoidance (rapid development, high gas exchange metabolism) and stress tolerance (slow development, low gas exchange metabolism), and that genetic variation in these character combinations may be maintained by environmental variation in stress levels in the species’ ruderal habitat. Received: 28 March 1996 / Accepted: 13 August 1996     

Carbon Isotope Discrimination in Coffee Genotypes Grown under Limited Water Supply

Photosynthetic gas exchange, plant-water relations characteristics, and stable carbon isotope discrimination (Δ) were evaluated for five Coffea arabica L. genotypes growing under two soil moisture regimes in the field. The Δ of leaf tissue was strongly correlated (r = −0.95) with inherent water use efficiency (ratio of assimilation to stomatal conductance; A/g). The variation in inherent water use efficiency (WUE) among genotypes was 30% for plants irrigated weekly. The higher WUE exhibited by some of these plants resulted from reduced g rather than increased photosynthetic capacity at a given g. Withholding irrigation for 1 month caused Δ to decline substantially in expanding leaf tissue of all genotypes. A strong correlation (r = 0.92) was found between Δ and plant hydraulic efficiency estimated as the ratio of g to the diurnal range in leaf water potential (Ψ_l). The Δ values for plants irrigated weekly adequately predicted drought-induced changes in Δ (r = 0.99) and midday Ψ_l (r = 0.95). The results indicated that Δ might be used to evaluate several aspects of plant performance and response to specific environmental conditions, once suitable background physiological data have been gathered.     

Variability in water use efficiency at the leaf level among Mediterranean plants with different growth forms

Assessing natural variability of leaf water use efficiency in plants adapted to extreme conditions of the Mediterranean climate represents an important step in the evaluation of the usefulness of some plant ecophysiological traits under water stress. Eleven Mediterranean species naturally inhabiting the Balearic Islands and corresponding to different growth forms (herbs, semi-deciduous shrubs, woody evergreen shrubs and woody evergreen semi-shrubs) were subject to progressive soil water depletion. Leaf intrinsic water use efficiency was measured by gas exchange at four different degrees of water stress. Under well watered conditions, differences in leaf intrinsic water use efficiency (A _N/g _s) among growth forms were limited to woody evergreen semi-shrubs, which presented the highest values. Under water stress conditions, differences became more evident, with a trend for an increase in A _N/g _s from woody evergreen shrubs, through semi-deciduous shrubs and herbaceous to woody evergreen semi-shrubs. The observed variation in A _N/g _s correlated with several physiological (leaf water potential, soil to leaf hydraulic conductance and stomatal conductance) and morphological (stomatal density) parameters, displaying a general relationship for all growth forms. This suggests that the capacity for withstanding water limitation is adaptive for all Mediterranean species. However, when A _N/g _s was related to leaf mass area, this relationship was not generally applicable, and depended on growth forms, suggesting that different growth forms display specific morphological adjustments in response to water shortage.     

Comparing the intersex genetic correlation for fitness across novel environments in the fruit fly,Drosophila serrata

Sexually antagonistic genetic variation can pose limits to the independent evolution and adaptation of the sexes. The extent of sexually antagonistic variation is reflected in the intersex genetic correlation for fitness (r_w^FM). Previous estimates of this correlation have been mostly limited to populations in environments to which they are already well adapted, making it difficult to gauge the importance of sexually antagonistic genetic variance during the early stages of adaptation, such as that occurring following abrupt environmental change or upon the colonization of new habitat. Here we assayed male and female lifetime fitness in a population of Drosophila serrata in four novel laboratory environments. We found that r_w^FM varied significantly across environments, with point estimates ranging from positive to negative values of considerable magnitude. We also found that the variability among estimates was because, at least in part, of significant differences among environments in the genetic variances of both male and female fitness, with no evidence of any significant changes in the intersex covariance itself, although standard errors of these estimates were large. Our results illustrate the unpredictable nature of r_w^FM in novel environments and suggest that, although sexually antagonistic genetic variance can be pronounced in some novel environments, it may have little effect in constraining the early stages of adaptation in others.     

Unthinned slow-growing ponderosa pine (Pinus ponderosa) trees contain muted isotopic signals in tree rings as compared to thinned trees

Key message

The muted wood isotopic signal in slow-growing trees of unthinned stands indicates lower responsiveness to changing environmental conditions compared to fast-growing trees in thinned stands.

Abstract

To examine the physiological processes associated with higher growth rates after thinning, we analyzed the oxygen isotopic values in wood (δ¹⁸O_w) of 12 ponderosa pine (Pinus ponderosa) trees from control, moderately, and heavily thinned stands and compared them with wood-based estimates of carbon isotope discrimination (?¹³C), basal area increment (BAI), and gas exchange. We found that (heavy) thinning led to shifts and increased inter-annual variability of both stable carbon and oxygen isotope ratios relative to the control throughout the first post-thinning decade. Results of a sensitivity analysis suggested that both an increase in stomatal conductance (g _s) and differences in source water among treatments are equally probable causes of the δ¹⁸O_w shift in heavily thinned stands. We modeled inter-annual changes in δ¹⁸O_w of trees from all treatments using environmental and physiological data and found that the significant increase in δ¹⁸O_w inter-annual variance was related to greater δ¹⁸O_w responsiveness to changing environmental conditions for trees in thinned stands when compared to control stands. Based on model results, the more muted climatic response of wood isotopes in slow-growing control trees is likely to be the consequence of reduced carbon sink strength causing a higher degree of mixing of previously stored and fresh assimilates when compared to faster-growing trees in thinned stands. Alternatively, the muted response of δ¹⁸O_w to climatic variation of trees in the control stand may result from little variation in the control stand in physiological processes (photosynthesis, transpiration) that are known to affect δ¹⁸O_w.     

The influence of ABA on water relation, photosynthesis parameters, and chlorophyll fluorescence under drought conditions in two maize hybrids with contrasting drought resistance

Drought is a major limitation of maize cultivation in Brazil. Agronomic and physiological practices have been considered to overcome this stress and consequently, increase grain production. The present study investigated the role of abscisic acid (ABA) application in some physiological parameters, in two hybrids with contrasting drought resistance (DKB 390 and BRS 1030 resistant and sensitive, respectively). Contrasting resistance to drought in these genotypes was determined in previous studies. Water deficit was imposed for 10 days at flowering stage, in association with the application of 100 μM abscisic acid on plant canopy. Evaluations of gas exchange, chlorophyll fluorescence, relative water content (RWC), and endogenous ABA content were performed during stress period and also at water recovery (recovery irrigation). A significant functional relationship was observed between RWC and the parameters of gas exchange and fluorescence. During water recovery, no differences were observed among the treatments. DKB 390 presented higher photosynthesis rate (P _n) and electron transport rate (ETR) under water stress, while BRS 1030 presented higher intercellular CO₂ concentration (C _i) and lower photochemical quenching (qP), non-photochemical quenching (NPQ), and lower F _v/F _m ratio. DBK 390 was more responsive to ABA application than BRS 1030, presenting higher endogenous ABA content in the first day of stress. DBK 390 with ABA application reduced the effect of water stress through maintenance of water status, an increase of photosynthetic parameters, and a decrease of decline in the functions of photosystem II during stress.     

Yield and morpho-agronomical evaluation of food-grade white sorghum hybrids grown in Southern Italy

Abstract

Grain sorghum [Sorghum bicolor (L.) Moench] is a gluten-free grain that is gaining attention as a food crop that can be used in the management of celiac disease. At present, sorghum is widely grown in many semiarid regions of the world. New food-grade sorghum cultivars are of particular interest in Mediterranean countries due to improved quality characteristics and gluten-free status of the grains. Until now very few studies have examined the grain yield (GYLD) and agronomic performance characteristics of food-grade sorghum hybrids in Italy. A 2 year study was conducted to evaluate the agronomic performance of eight food-grade sorghum hybrids representing different maturity classes in trials conducted in Southern Italy. The results showed wide variation in adaptation of these hybrids as measured by differences in GYLD (2.35–8.50 t ha^?1) and other pheno-morphological characteristics. Of particular interest was the fact that many of the early-flowering hybrids (e.g. SP-X303) performed better than the later-flowering hybrids (e.g. ArchX-02). These results demonstrated that flowering time of hybrid and crop cycle length are important factors to consider in selecting cultivars for production in the Mediterranean region.     

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.