Osmotic adjustment in sorghum: I. Mechanisms of diurnal osmotic potential changes.

Osmotic adjustment, defined as a lowering of osmotic potential (psi(pi)) due to net solute accumulation in response to water stress, has been considered to be a beneficial drought tolerance mechanism in some crop species. The objective of this experiment was to determine the relative contribution of passive versus active mechanisms involved in diurnal psi(pi) changes in sorghum (Sorghum bicolor L. Moench) leaf tissue in response to water stress. A single sorghum hybrid (cv ATx623 x RTx430) was grown in the field under variable water supplies. Water potential, psi(pi), and relative water content were measured diurnally on expanding and the uppermost fully expanded leaves before flowering and on fully expanded leaves during the grain-filling period. Diurnal changes in total osmotic potential (Deltapsi(pi)) in response to water stress was 1.1 megapascals before flowering and 1.4 megapascals during grain filling in comparison with 0.53 megapascal under well-watered conditions. Under water-stressed conditions, passive concentration of solutes associated with dehydration accounted for 50% (0.55 megapascal) of the diurnal Deltapsi(pi) before flowering and 47% (0.66 megapascal) of the change during grain filling. Net solute accumulation accounted for 42% (0.46 megapascal) of the diurnal Deltapsi(pi) before flowering and 45% (0.63 megapascal) of the change during grain filling in water-stressed leaves. The relative contribution of changes in nonosmotic volume (decreased turgid weight/dry weight) to diurnal Deltapsi(pi) was less than 8% at either growth stages. Water stress did not affect leaf tissue elasticity or partitioning of water between the symplasm and apoplasm.

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 (Psi(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 ATx623 x RTx430) was studied under field conditions using four different water supplies. Diurnal and midday water potential, solute potential, Psi(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 (BTx623 and BTx378) 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, BTx623 maintained significantly higher g and A than BTx378 during the water stress cycle. However, no significant differences in OA or Psi(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.

Leaf photosynthetic rate is correlated with biomass and grain production in grain sorghum lines.

Significant genetic variation in leaf photosynthetic rate has been reported in grain sorghum [Sorghum biocolor (L.) Moench]. The relationships between leaf photosynthetic rates and total biomass production and grain yield remain to be established and formed the purpose of this experiment. Twenty two grain sorghum parent lines were tested in the field during the 1988 growing season under well-watered and water-limited conditions. Net carbon assimilation rates were measured at mid-day during the 30 day period from panicle initiation to head exertion on upper-most fully expanded leaves using a portable photosynthesis system (LI-6200). Total biomass and grain production were determined at physiological maturity. The lines exhibited significant genetic variation in leaf photosynthetic rate, total biomass production and grain yield. Significant positive correlations existed between leaf photosynthesis and total biomass and grain production under both well-watered and water-limited conditions. The results suggest that leaf photosynthetic rate measured prior to flowering is a good indicator of productivity in grain sorghum.

Genetic variation for gas exchange rates in grain sorghum.

Carbon assimilation rate (A) and stomatal conductance (g) are highly correlated. However, the slope of the A versus g relationship differs among species and environments resulting in differences in gas exchange efficiency which should reflect water use efficiency. The objective of this research was to determine the genetic variation for A and g in grain sorghum (Sorghum bicolor [L.] Moench.). Field experiments were conducted using 30 sorghum hybrids with four water supply treatments. A, g, and leaf water potential (Psi(w)) of individual leaves were monitored every 15 to 20 days. Significant genetic variation existed among the hybrids for A and g. Plant age and water supply also affected A and g as expected. When A was regressed on g for each hybrid, large and significant differences existed among the slopes, implying differences in intrinsic gas exchange efficiency. The regression analysis of A and g versus Psi(w) suggested that A was more sensitive than g to increasing water stress. Genetic differences in the rate of change in A as water stress increased were observed. Regression analysis was used to evaluate the individual hybrid response relative to other hybrids. Twofold difference in slopes existed for A. These results provide evidence for genetic variation in gas exchange rates which might directly contribute to whole plant water use efficiency and productivity.

Photosynthetic rate control in cotton : stomatal and nonstomatal factors.

The relationship between single leaf photosynthesis and conductance was examined in cotton (Gossypium hirsutum L.) across a range of environmental conditions. The purpose of this research was to separate and define the degree of stomatal and nonstomatal limitations in the photosynthetic process of field-grown cotton.Photosynthetic rates were related to leaf conductance of upper canopy leaves in a curvilinear manner. Increases in leaf conductance of CO(2) in excess of 0.3 to 0.4 mole per square meter per second did not result in significant increases in gross or net photosynthetic rates. No tight coupling between environmental influences on photosynthetic rates and those affecting conductance levels was evident, since photosynthesis per unit leaf conductance did not remain constant. Slowly developing water stress caused greater reductions in photosynthesis than in leaf conductance, indicating nonstomatal limitations of photosynthesis.Increases in external CO(2) concentration to levels above ambient did not produce proportional increases in photosynthesis even though substomatal or intercellular CO(2) concentration increased. The lack of a linear increase in photosynthetic rate in response to increases in leaf conductance and in response to increases in external CO(2) concentration demonstrated that nonstomatal factors are major photosynthetic rate determinants of cotton under field conditions.

Photosynthetic rate control in cotton : photorespiration.

The purpose of this research was to determine the magnitude of photorespiration in field-grown cotton (Gossypium hirsutum L.) as a function of environmental and plant-related factors. Photorespiration rates were estimated as the difference between measured gross and net photosynthetic rates.A linear increase in photorespiration was observed as air temperature increased from 22 to 40 degrees C at saturating photon flux density. At 22 degrees C, photorespiration was less than 15 per cent of net photosynthesis and very comparable to the dark respiration rate. At 40 degrees C, photorespiration represented about 50 per cent of net photosynthesis. Gross photosynthesis had a temperature optimum of 32 to 34 degrees C. Water stress, as indicated by Psi(L), did not alter the ratio of gross photosynthesis to net photosynthesis when the confounding effects of leaf temperature differences were accounted for in the data analyses. A reduction in both gross and net photosynthesis was apparent as Psi(L) declined from -2.0 megapascals indicating direct effects of water stress on the photosynthetic process. Photorespiration expressed as a proportion of net photosynthesis increased as water stress intensified.Cotton cultivars possessing a fruit load had significantly higher gross and net photosynthetic rates and lower photorespiration rates than did photoperiod-sensitive cotton strains without a fruit load. Within the fruiting types, which were genetically very similar, only minor differences were observed in the photorespiration:net photosynthesis ratios. However, in the photoperiod-sensitive strains, considerable genetic variability existed when photorespiration was expressed as a proportion of net photosynthesis. These results suggest that the kinetics of ribulose-1,5-bisphosphate carboxylase:oxygenase may be different and, thus, the possibility of genetically reducing photorespiration exists.

Relative sensitivity of photosynthetic assimilation and translocation of carbon to water stress.

The relationship between photosynthesis and translocation rate changes as affected by water stress intensity and stage of plant development was evaluated in cotton and sorghum, representing a C(3) and a C(4) photosynthetic type, respectively. Photosynthetic rates were reduced as midday leaf water potentials declined from -14 to -27 bars in both species. Sorghum maintained higher photosynthesis and translocation rates compared to cotton at comparable leaf water potentials; however, the rate of change per bar decline in water potential was greater in sorghum than in cotton. Photosynthetic rates were reduced with increasing water stress prior to any significant change in translocation rates suggesting that photosynthesis is the more sensitive of the two processes. Severe water stress, corresponding to leaf water potentials of -27 bars, did not completely inhibit either photosynthesis or translocation.

Stomatal and nonstomatal regulation of water use in cotton, corn, and sorghum.

Stomata of corn (Zea mays L.) and sorghum (Sorghum bicolor L.) responded to changes in leaf water potential during the vegetative growth phase. During reproductive growth, leaf resistances were minimal and stomata were no longer sensitive to bulk leaf water status even when leaf water potentials approached -27 bars. Stomata of corn, cotton (Gossypium hirsutum L.), and sorghum appear to respond to changes in the humidity deficit between the leaf and air and in this manner, regulated transpirational flux to some degree. Distinct differences in water transport efficiency were observed in the three species. Under nonlimiting soil water conditions, sorghum exhibited the greatest efficiency of water transport while under limiting soil moisture conditions, cotton appeared most efficient. Corn was the least efficient with respect to nonstomatal regulation of water use. Differences in drought tolerance among the three species are partially dependent on stomatal regulation of water loss, but efficiency of the water transport system may be more related to drought adaptation. This is particularly important since stomata of all three species did not respond to bulk leaf water status during a large portion of the growing season.