Inheritance of osmotic adjustment to water stress in three grain sorghum crosses.

Water stress is one of the major constraints to the grain yield of sorghum in tropical and sub-tropical areas of the world. Osmotic adjustment has been widely proposed as a plant attribute that confers adaptation to water stress. The inheritance of osmotic adjustment to water stress was investigated in a series of generations derived from the three possible bi-parental crosses between two inbred sorghum lines with a high capacity for osmotic adjustment (Tx2813 and TAM422; high-OA lines) and one with a low capacity (QL27; low-OA line). Broad-sense heritability on a single-plant basis was generally found to be high. Analysis of segregation ratios by the mixture method of clustering identified two independent major genes for high osmotic adjustment. The line Tx2813 possessed a recessive gene which is given the symbol oa1; the line TAM422 possessed an additive gene which is given the symbol OA2. There was some evidence that there may be other minor genes which influence the expression of osmotic adjustment in these crosses as two putative transgressive segregants, with higher osmotic adjustment than the parents, were identified from the cross between Tx2813 and TAM422. Populations of recombinant inbred lines were developed and characterised for osmotic adjustment for two of the crosses (QL27 x TAM422, low-OA x high-OA; Tx2813 x TAM422, high-oal x high-OA2). These will be used to conduct experiments which test hypotheses about the contribution of the high-osmotic-adjustment genes to the grain yield of sorghum under a range of water-stress conditions.

DNA polymorphisms in grain sorghum (Sorghum bicolor (L.) Moench).

Molecular markers [random amplified polymorphic DNA (RAPD) and restriction fragment length polymorphism (RFLP)] were used to determine the frequency of DNA polymorphism in grain sorghum (Sorghum bicolor (L.) Moench). Twenty-nine oligonucleotide primers were employed for RAPDs, generating a total of 262 DNA fragments, of which 145 were polymorphic in at least one pairwise comparison between 36 genotypes. Individual primers differed significantly in their ability to detect genetic polymorphism in the species. The overall frequency of polymorphisms was low with a mean frequency of 0.117 polymorphisms per RAPD band being obtained from all pairwise comparisons between genotypes, with maximum and minimum values of 0.212 and 0.039, respectively. Results from phenetic analysis of bandsharing data were consistent with current sub-specific groupings of the species, with clusters of Durra, Zerazera, Caud-Nig, Caud-Kaura and Caffrorum being discernible. The results also indicated that individuals of a similar taxonomic grouping but different geographic origin may be genetically less identical than previously considered. Similar frequencies of polymorphism to that obtained with RAPDs were obtained with RFLPs. Results from these experiments indicated that a high level of genetic uniformity exists within S. bicolor.

Comparative responses of the Savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation.

Two perennial tussock grasses of savannas were compared in a glasshouse study to determine why they differed in their ability to withstand frequent, heavy grazing; Cenchrus ciliaris is tolerant and Themeda triandra is intolerant of heavy grazing. Frequent defoliation at weekly intervals for six weeks reduced shoot biomass production over a subsequent 42 day regrowth period compared with previously undefoliated plants (infrequent) in T. triandra, but not in C. ciliaris. Leaf area of T. triandra expanded rapidly following defoliation but high initial relative growth rates of shoots were not sustained after 14 days of regrowth because of reducing light utilising efficiency of leaves. Frequently defoliated plants were slower in rate of leaf area expansion and this was associated with reduced photosynthetic capacity of newly formed leaves, lower allocation of photosynthate to leaves but not lower tiller numbers. T. triandra appears well adapted to a regime where defoliation is sufficiently infrequent to allow carbon to be fixed to replace that used in initial leaf area expansion. In contrast, C. ciliaris is better adapted to frequent defoliation than is T. triandra, because horizontally orientated nodal tillers are produced below the defoliation level. This morphological adaptation resulted in a 10-fold higher leaf area remaining after defoliation compared with similarly defoliated T. triandra, which together with the maintenance of moderate levels of light utilising efficiency, contributed to the higher leaf area and shoot weight throughout the regrowth period.

Photosynthetic responses of native and introduced C4 grasses from Venezuelan savannas.

Introduced African grasses are invading the grasslands of the Venezuelan savannas and displacing the native grasses. This work, which is part of a program to understand the reasons for the success of the African grasses, specifically investigates whether introduced and native grasses differ in some photosynthetic characteristics.The responses to photon flux density, leaf temperature, leaf-air vapour pressure difference and leaf water potential of leaf photosynthetic rate of two introduced African C4 grasses (Hyparrhenia rufa and Melinis minutiflora) and of a lowland and a highland population of a native Venezuelan grass (Trachypogon plumosus) grown under controlled conditions were compared. These responses in all three species were typical of tropical C4 pasture grasses. The introduced grasses had higher maximum leaf conductance, net photosynthetic rates, and optimum temperature (H. rufa only) for photosynthesis than T. plumosus. However, T. plumosus was able to continue photosynthesis to lower leaf water potentials than the two introduced grasses, and the efficiency which it utilized water, light and mineral nutrients to fix carbon were similar to those of the introduced grasses.The higher rates of leaf photosynthesis of the introduced grasses contributed to, but only partially explained, the higher growth rates compared to T. plumosus. The higher growth rates and nutrient concentration of the introduced grasses are consistent with their ability to establish rapidly, compete successfully for resources, and displace T. plumosus from moist, fertile sites. Conversely, the slower growth rate, lower nutrient concentrations, and superior water relations characteristics are consistent with the capacity of T. plumosus to resist invasion by introduced grasses in poorer sites.

Paraheliotropic leaf movement in Siratro as a protective mechanism against drought-induced damage to primary photosynthetic reactions: damage by excessive light and heat.

Damage to primary photosynthetic reactions by drought, excess light and heat in leaves of Macroptilium atropurpureum Dc. cv. Siratro was assessed by measurements of chlorophyll fluorescence emission kinetics at 77 K (-196°C). Paraheliotropic leaf movement protected waterstressed Siratro leaves from damage by excess light (photoinhibition), by heat, and by the interactive effects of excess light and high leaf temperatures. When the leaves were restrained to a horizontal position, photoinhibition occurred and the degree of photoinhibitory damage increased with the time of exposure to high levels of solar radiation. Severe inhibition was followed by leaf death, but leaves gradually recovered from moderate damage. This drought-induced photoinhibitory damage seemed more closely related to low leaf water potential than to low leaf conductance. Exposure to leaf temperatures above 42°C caused damage to the photosynthetic system even in the dark and leaves died at 48°C. Between 42 and 48°C the degree of heat damage increased with the time of exposure, but recovery from moderate heat damage occurred over several days. The threshold temperature for direct heat damage increased with the growth temperature regime, but was unaffected by water-stress history or by current leaf water status. No direct heat damage occurred below 42°C, but in water-stressed plants photoinhibition increased with increasing leaf temperature in the range 31-42°C and with increasing photon flux density up to full sunglight values. Thus, water stress evidently predisposes the photosynthetic system to photoinhibition and high leaf temperature exacerbates this photoinhibitory damage. It seems probable that, under the climatic conditions where Siratro occurs in nature, but in the absence of paraheliotropic leaf movement, photoinhibitory damage would occur more frequently during drought than would direct heat damage.

The distribution of leaf photosynthetic activity in a mixed grass-legume pasture canopy.

The distribution of net photosynthetic activity of leaves was measured in a mixed grass (Setaria sphacelata var. sericea)-legume (Desmodium intortum) pasture stand using a method based on concurrent measurement of the rate of CO2 exchange, and (14)CO2 dosing followed by rapid harvesting according to height strata. Comparisons were also made between plots which differed in the period of regrowth following defoliation.The usual superiority of leaf net photosynthetic rates of a C4 grass, compared with C3 legume leaves, was found in the upper, well illuminated strata. These rates were, however, much lower than those usually described for horizontally exposed leaves, primarily because leaves in the pasture stand were inclined to the horizontal. At greater depth in the canopies, the superiority of rates in the grass was less evident, and consequently the relative contributions of grass and legume to canopy photosynthesis became more dependent on their leaf area indices.Attention is drawn to the relative simplicity of the method for examining the contribution of leaves, which may differ according to species or position in the canopy, to productivity of the whole stand.

The distribution of leaf photosynthetic activity in a mixed grass-legume pasture canopy.

The distribution of net photosynthetic activity of leaves was measured in a mixed grass (Setaria sphacelata var. sericea)-legume (Desmodium intortum) pasture stand using a method based on concurrent measurement of the rate of CO2 exchange, and (14)CO2 dosing followed by rapid harvesting according to height strata. Comparisons were also made between plots which differed in the period of regrowth following defoliation.The usual superiority of leaf net photosynthetic rates of a C4 grass, compared with C3 legume leaves, was found in the upper, well illuminated strata. These rates were, however, much lower than those usually described for horizontally exposed leaves, primarily because leaves in the pasture stand were inclined to the horizontal. At greater depth in the canopies, the superiority of rates in the grass was less evident, and consequently the relative contributions of grass and legume to canopy photosynthesis became more dependent on their leaf area indices.Attention is drawn to the relative simplicity of the method for examining the contribution of leaves, which may differ according to species or position in the canopy, to productivity of the whole stand.

Effect of temperature on light utilization efficiency of leaves in C3 legumes and C 4 grasses.

The effect of temperatures between 15 and 50°C on the light utilization efficiency of leaf net photosynthesis was studied in two C4 grasses and two C3 legumes. In the legumes, light utilization efficiency (µ mol of CO2 fixed per µ Einstein of incident photosynthetic quantum flux) declined linearly with temperature between 15 and 45°C. On the other hand, it was independent of temperature between 15 and 40°C in the grasses, but it declined rapidly between 40 and 50°C. Values for grasses were greater than those for legumes at all temperatures; for example, at 30°C the ratio of the mean values of grass to legume was 1.6-1.7 which is similar to a previously published value of 1.6. Light utilization efficiency was similar for grasses and legumes only at temperatures below 10 and above 50°C. These findings are compared with published data, especially those of Ehleringer and Björkman, and their implications for predicting or interpreting the distribution of C3 and C4 species are briefly discussed.

Stomatal responses to humidity in Opuntia inermis in relation to control of CO2 and H2O exchange patterns.

At constant cladode temperature the stomatal resistance of O. inermis increased when the cladode-air vapor pressure difference was increased and stomatal resistance decreased when the cladode-air vapor pressure difference was lowered. Net CO2 fixation in the dark was very responsive to these humidity dependent changes in stomatal resistance. Net CO2 fixation and stomatal resistance in the light did not respond to changes in cladode-air vapor pressure differences in the light under the conditions tested. When temperature response functions for dark CO2 fixation were examined at constant ambient humidity, the reduction in dark CO2 fixation at higher temperatures was largely due to stomatal closure in response to the increased vapor pressure difference. The water requirement for net CO2 fixation in the dark at typical nocturnal vapor pressure differences was about 10 times lower than that of net CO2 fixation in the light at vapor pressure differences typical of the late afternoon. The role of the stomatal responses to humidity in determining the patterns and rates of net CO2 exchange in the light or dark, and its possible ecological significance is discussed.

Comparison between pressure-volume and dewpoint-hygrometry techniques for determining the water relations characteristics of grass and legume leaves.

The water relations characteristics of three grass species (Panicum maximum var. trichoglume, Cenchrus ciliaris, Heteropogon contortus), and a legume (Macroptilium atropurpureum) grown in the field were measured using both a modified pressure/volume technique with pressure bomb measurements on single leaves and a dewpoint hygrometry technique applied to fresh and to frozen and thawed leaf discs.The two techniques agreed well in the estimates of osmotic potential at full turgor and the water potential at zero turgor. However, for parameters such as the relative water content at zero turgor, bound water and bulk modulus of elasticity there was a poor correlation between the estimates from the two methods. The pressure/volume technique gave less variable results and is more convenient for field use than the hygrometry technique. The determination of the modulus of elasticity from various functions relating pressure potential to relative water content is discussed.

Effect of oxygen concentration on leaf photosynthesis and resistances to carbon dioxide diffusion.

Net photosynthesis of tropical legume leaves increased by 44% and that of tropical grass leaves was unaffected when oxygen concentration was reduced from 21 to 0.2%. Stomatal resistance to carbon dioxide diffusion was unaltered in both cases but mesophyll resistance of legume leaves decreased with oxygen concentration. It is proposed that the decrease in mesophyll resistance is accompanied by decreases in excitation and carboxylation resistances.