Search

Two grasses and two legumes were grown at three illuminances: one grass and one legume in a growth cabinet (100, 50 and 34 % relative illuminance) and one of each in a glasshouse (100, 30 and 10% relative illuminance). The response to shading was investigated by using classical growth analysis techniques, and the photosynthesis-respiration balance obtained by Watson and Hayashi's method. Shading caused a reduction in the proportion of root and a corresponding increase in the proportion of leaf, and an increase in the shoot/root ratio in all species except green panic. The relative growth rates of grasses (which were only clearly superior at the highest illuminance) were affected more by shading than were those of legumes, and the greater shading effect resulted from a greater decrease in net assimilation rate, and to a lesser extent, from a smaller compensatory increase in leaf area ratio in the grasses than in the legumes. The greater sensitivity of net assimilation rate of grasses to shading arose from a stronger dependence of the photosynthetic rate on illuminance. The significance of these findings for pasture production is discussed. Both tiller and runner production were dependent upon the total amount of radiant energy received by the plant, and in the growth cabinet, at least, were independent of its intensity, duration and sequence.

A comparison is made between two models of canopy photosynthesis for monocultures that differ in their treatment of the variation of the rate of single-leaf photosynthesis in response to the irradiance in which the leaves have grown. Both are shown to be deficient, and a simple modification to one that results in a much improved performance is presented. The theory uses the non-rectangular hyperbola for the rate of single-leaf photosynthesis and incorporates the Monsi-Saeki approach for the light-intercepting characteristics of the canopy. The model is then extended to describe the photosynthesis of mixed canopies, which requires knowledge of the relative position of each leaf-area component within the sward. Analytical expressions can be derived for the case where the leaf-area components are homogeneously distributed relative to each other through the depth of the canopy. This is shown to be a good representation of leaf-area distributions in continuously grazed temperate grass-clover swards as well as frequently cut tropical grass-legume swards, and so is an important special case.

This work was undertaken to determine if the stimulation of growth associated with shading of some tropical C4 grasses growing on soils with low to moderate nitrogen availability is partly due to overcoming photoinhibition (i. e. damage caused by excessive light). Four grasses (green panic, carpet grass, buffalo grass and kikuyu) were grown in full sunlight and at 37% of full sunlight, and given a low or high nitrogen supply. Despite differences of up to twofold in leaf nitrogen and chlorophyll concentration between high and low nitrogen treatments in all four grasses, photoinhibition measured by reduction in chlorophyll fluorescence was less than 3% in leaves of low nitrogen content that developed in full sunlight. Therefore, photoinhibition is not a contributor to the poor growth of nitrogen-limited grasses in full sunlight. A second objective was to determine if low nitrogen content predisposed shade-grown leaves to photoinhibition when they were subsequently exposed to full sunlight. Green panic plants that had been given either high or low nitrogen supply and grown in 37% of full sunlight were transferred to full sunlight and the extent of photoinhibition was followed over 8 days. The amount of photoinhibition that occurred was small (<6%) compared with plants grown at either nitrogen level in full sunlight. Therefore, shade and low nitrogen content separately or in combination did not cause any appreciable photoinhibition in green panic.

The significance of photoinhibition induced by water stress for growth and grain yield of sorghum was investigated by shading water-stressed plants for a 7-day period during panicle development. Thereafter, shading was removed, stressed plants were rewatered, and all plants were kept well-watered until physiological maturity. Shading water-stressed plants for a 7-day period increased grain number and grain yield of primary heads by 10- and 13-fold, respectively. Even though plants were severely stressed, photoinhibition was only 20%. Moreover, reduction of photoinhibition by shading did not improve carbon supply to the developing panicle. Therefore, the response of grain yield to shading was attributed to a 15% higher average leaf water potential in shaded plants, which arose despite attempts to keep water status similar in shaded and unshaded plants. The effect of water stress on the primary head was so severe that many plants produced secondary heads. The grain yield from these secondary heads compensated for the low yield of primary heads so that the total dry matter production and total grain yield of water-stressed plants were similar to those of unstressed plants. In view of the ability of this relatively determinate species to compensate for potential yield losses, as well as the mechanisms to avoid water stress, high temperatures and excessive absorption of light, it seems unlikely that photoinhibition induced by water stress will be of agronomic significance in grain sorghum.

The water balance of soybean (Glycine max), cowpea (Vigna unguiculata), black gram (Vigna mungo), and pigeonpea (Cajanus cajan) grown in pots was studied during a soil drying cycle. The response of the plants was analysed for three distinct stages of dehydration. In stage I, the rate of transpiration remained constant and equal to that of well watered plants even though soil water status fell by more than 50%. Stage II began when the rate of soil water supply to the plant was less than potential transpiration and stomates closed resultingjn the maintenance of plant water balance. When soil water content was expressed as a fraction of transpirable soil water, all species showed a transition from stage I to stage II at a fraction of transpirable soil water of about 0.3 to 0.2. As the soil water declined further, all species had a similar decrease in relative transpiration rate. Consequently, the responses of the four species in stages I and II were essentially identical, except that pigeonpea extracted a slightly greater amount of soil water. Stage III occurred once stomates had reached minimum conductance and water loss was then a function of the epidermal conductance and the environment around the leaf. Substantial differences were found among the four grain legumes in epidermal conductance. Soybean had the highest conductance, followed by black gram, cowpea and pigeonpea. Substantial variation in dehydration tolerance among the four grain legumes was also found: the ranking of dehydration tolerance based on the relative water content was pigeonpea > cowpea > mungbean > soybean. Differences among the four grain legume species in the duration of stage III which finished when plants died, were consistent with differences in epidermal conductance and in dehydration tolerance of leaves.

The nature of C3 and C4 pasture plants is described, and the potential biomass productivity is discussed. Then differences in net annual production and crop growth rates of C3 and C4 pasture plants are analysed in terms of the interception of solar radiation and the efficiency of its conversion to dry matter. Finally, the components of photosynthetic production of tropical C4 grasses and tropical C3 legumes are compared with their growth rates and dry matter yields in the subtropics. The higher crop growth rates of tropical C4 grasses compared with C3 pasture plants, whether they are of temperate or tropical origin, seem to be due mainly to higher leaf net photosynthetic rates and to the absence of photorespiration associated with possession of the C4 pathway. However, light utilization efficiencies are probably also higher in C4 grasses. In addition to these two characteristics, the longer growing season of C4 grasses in the warm conditions of the tropics contributes to their higher net annual production of dry matter. There is a close association between possession of the C4 pathway, higher rates of photosynthesis and growth, and higher dry matter yields when tropical C4 pasture grasses are compared with tropical C3 legumes in a subtropical climate.

Diaheliotropic movements in Macroptilium atropurpureum cv. Siratro caused the adaxial surfaces of leaflets to orient towards a light by the flexing of secondary pulvini at the base of the leaflets, and to a lesser extent by petiole bending and flexing of the primary pulvinus at the base of the petiole. Secondary pulvini flexed more in response to light than did primary pulvini. This flexing was in response to blue light such that the illuminated side of the pulvinus contracted, and the leaflet reoriented with its adaxial surface facing the light. This response was enhanced by the following: a light gradient along the length of the illuminated surface of the pulvinus, the presence of additional low-intensity diffuse light, a photon irradiance greater than 0.5 mmol m-2 s-1, and a change of photon irradiance. The abaxial surface of a pulvinus was more sensitive to light than was its adaxial surface. The stimulus for movement was transmitted from an illuminated primary pulvinus to the shaded secondary pulvini of the same leaf, and this caused re-orientation of both individual leaflets and the whole leaf. Similarly, a stimulus was transmitted to the primary pulvinus from secondary pulvini (or from the adjacent petiole) when illuminated, resulting in a deflection of the petiole by the primary pulvinus and re-orientation of the adaxial surface of the whole leaf towards the light. Secondary pulvini have the ability to move individual leaflets in three dimensions because the plane of the leaflet is at right angles, in two dimensions, to the axis of the distal end of the pulvinus. Terminal leaflets showed a greater three-dimensional movement than did lateral leaflets. Movement of a light through large angles resulted in leaflet re-orientation proportional to, but less than, the change in angle of the light. On the other hand, secondary pulvini caused leaflets to track, though imperfectly, a light source that simulated the movement of the sun. A circumnutational circadian rhythm is partly responsible for the diaheliotropic movements of Siratro leaves outdoors, and the ability of an individual leaflet to track the sun depends upon its orientation with respect to the sun.

Stomatal conductance (g) and leaf photosynthetic rate (P) of many species in the field are often less sensitive to water deficits than when grown in small pots under controlled conditions. This may result from stomatal adjustment in field-grown plants in response to water deficits that develop slowly, whereas adjustment does not occur under the rapid drying experienced by plants in small pots. To test this hypothesis we studied the response to water potential (Ψl) of g and P in three tropical C4 grasses, green panic (Panicum maximum var. trichoglume), buffel grass (Cenchrus ciliaris) and spear grass (Heteropogon contortus) and a tropical legume, Siratro (Macroptilium atropurpureum), grown under controlled conditions and in the field. Field experiments clearly showed that stomatal adjustment occurred so that g and P were progressively less sensitive to the decline in Ψl as water deficits increased during a long soil drying cycle. For example in one experiment, the Ψl at which P approached zero fell from - 1.9, -2.0 and -2.4 MPa to -4.0, -4.0 and -3.3 MPa for green panic, spear grass and buffel grass, respectively. This stomatal adjustment was reversed within 10 days after rewatering to the well watered condition. Little stomatal adjustment occurred in plants grown under controlled conditions in small pots in which both soil water and Ψl fell rapidly. However, if plants were grown in similar conditions but in large pots of soil so that soil water and Ψl decreased slowly, stomatal adjustment comparable with field-grown plants was observed. Siratro showed much less stomatal adjustment than the grasses and Ψl at which P approached zero only fell from - 1.2 to - 1-5 MPa.

M. atropurpureum cv. Siratro and C. ciliaris cv. Biloela (buffel grass) plants in well-watered and droughted mixed swards were studied to investigate how the two species survive during periods of drought. Rates of photosynthesis and water use efficiencies were generally highest in well-watered buffel grass, whereas the conductance of the abaxial surfaces of leaves was highest in well-watered Siratro. The minimum and daily mean water potentials were consistently lower in well-watered buffel grass than in well-watered Siratro, indicating a lower hydraulic conductance in the soil-plant system of buffel grass. Gross photosynthesis and the conductance of the abaxial surfaces of leaves of both species fell with falling water potential, as did the water use efficiency of buffel grass. Droughted Siratro exhibited paraheliotropic leaf movement, and a reduction in stomatal aperture in the middle of the day. Despite its greater sensitivity to water deficits, Siratro remained physiologically active throughout at least a portion of the day even on the last day of a drying cycle when its leaf water potential ranged from -1 MPa at dawn to a minimum of -1.8 MPa, when the corresponding water potentials for buffel grass were about -4 MPa and about -5 MPa. Siratro had a higher water potential mainly because of the deeper penetration of its roots and its consequent access to water lower in the profile. However, the higher water potential was also due to stomatal opening, and therefore photosynthesis, in droughted Siratro occurring only at times of the day when evaporation, temperature and solar radiation were low. The conductance of the abaxial surfaces of droughted buffel grass leaves was generally steady, or declined gradually during the day. Buffel grass was able to survive drought because of its inherent desiccation tolerance, whereas Siratro was able to survive drought in a mixed sward because it did not compete with buffel grass for soil water.

Osmotic adjustment, calculated from the difference in concentrations of ions and sugars between stressed and unstressed leaves of green panic, buffel grass and speargrass given normal or high potassium supply in the field, was determined during a 37-day drying cycle and following rewatering. Pressure/volume (P/V) curves were used to characterize the water relations of buffel grass with normal K supply on eight occasions during the experiment, and of the remaining treatments only at the end of the drying cycle. The maximum osmotic adjustment for the three grasses was 0.84-1.0 MPa calculated from chemical analyses of solutes and 0.5-0.7 MPa derived from the P/V analyses. High K supply did not enhance osmotic adjustment of water-stressed leaves. Osmotic adjustment reached a plateau 22 days after watering ceased, and its relationship with leaf water potential (Ψ) was the same for all species. When midafternoon Ψ was c. -4.8 MPa and leaves were permanently wilted, osmotic adjustment was 90% of maximum, suggesting that osmotic adjustment ceased when growth and most physiological activity were curtailed by loss of turgor. Osmotic adjustment delayed the onset of permanent wilting in water-stressed leaves by c. 4 days. Therefore, its contribution to the maintenance of leaf growth during water stress in these three grasses was small.

The responses to water stress of five accessions representing four species of the legume Centrosema from contrasting moisture environments were compared under controlled conditions with those of Macroptilium atropurpureum cv. Siratro, a species which avoids dehydration. Species of Centrosema were able to tolerate leaf water potentials as low as -8 to -12 MPa, and all showed osmotic and stomatal adjustment. However, they differed in the tolerance of their leaves to water stress and in the leaf water potential at which stomata were effectively closed. There was a strong positive relationship between water stress tolerance of leaves and the leaf water potential for effective stomatal closure, among the Centrosema accessions and Siratro. The results are consistent with the natural ecological distribution of the species and their behaviour in different moisture environments in northern and north-eastern Australia.

Buffel grass was subjected to a soil drying cycle for 5 weeks in a semi-arid environment. As water stress developed, the leaf water relations characteristics of these plants (Dry treatment) were compared with those of irrigated plants (Wet treatment). Leaf water potential (Ψ) of the Dry treatment measured at 1400 h decreased to a minimum of -6.9 MPa. The stressed leaves adjusted osmotically, with the osmotic potential at full turgor (Ψπ100) decreasing (becoming more negative) linearly with time (0.017 MPa day-1) and with decreasing water potential measured at 1400 h (0.11 MPa per 1 MPa decrease in Ψ). Maximum osmotic adjustment (Ψπ100 Wet -Ψπ100 Dry) was 0.66 MPa, and this change together with lower cell wall elasticity decreased by 1.03 MPa the water potential (Ψ0) at which the stressed leaves lost turgor. Differences between the stress- acclimated Dry leaves and the Wet leaves in bound water, turgid weight:dry weight ratio and the relative water content at which they reached zero turgor were small and inconsistent. At 18 days after rewatering, the Ψπ100 value of acclimated leaves was still 0.18 MPa lower than that of the control leaves. The substantial shift in Ψ0 gained the stress-acclimated leaves only one extra day before they lost turgor at 1400 h, and only 2.5 extra days before being permanently wilted. This small gain in time and the rapid cessation of leaf growth even before positive turgor was completely lost suggests that osmotic adjustment may not contribute greatly to continued leaf growth in water-stressed plants of buffel grass.

The response was measured of stomatal conductance and leaf photosynthesis to changing leaf water potential in the legume siratro subjected to a sequence of I-week cycles of increasing soil water deficit followed by watering. The response of stomatal conductance was described using a continuous mathematical function, which is more robust and accurate than the usual discontinuous linear function used to analyse such data. After seven successive cycles of water deficit, there was no apparent adjustment of the short-term response of leaf conductance to leaf water potential.

The influence of leaf water potential (Ψl), saturation deficit (δe), and temperature on stornatal conductance (gs) of kenaf and sorghum was determined at non-limiting quantum fluxes both at high and low soil water status in the field, and under controlled environment conditions. Sorghum stomata closed in response to Ψl in a manner consistent with the classical feedback control, while kenaf stomata responded directly to δe, independently of Ψl. The rate of decrease of gs of kenaf with increasing δe was greater at higher temperatures. Also the stomata of stressed kenaf were less sensitive to δe than those of the well-watered crop due to the limitation in degree of stomatal opening imposed by water stress. The degree of stomatal adjustment in kenaf was also assessed. In the stressed kenaf, the Ψl at half the maximum gs was c. 0.17 MPa lower and gs was more sensitive to decreases in Ψl, than in the well watered control. However, there was little difference in the Ψl at 95% stomatal closure.

The extent of night-time stomatal opening in field-grown kenaf (Hibiscus cannabinus L.), a C3 dicotyledon, and sorghum [Sorghum bicolor (L.) Moench], a C4 grass, and the factors controlling the opening were studied at high and low soil water status. Since saturation deficit (δe) and temperature varied together in the field, the response of stomatal conductance (gs) to these individual factors was determined under controlled environment conditions in a leaf chamber apparatus. At both high and low soil water status, the stomata of sorghum were closed from sunset to sunrise, whereas with kenaf partial stomatal opening was observed throughout the night. Initiation of night opening occurred in response to decreasing temperature, but the degree of opening was determined by plant water status. The importance of night-time stomatal opening on the water relations of the crop is discussed.

Net photosynthesis of the last fully expanded leaf of P. maximum var. trichoglume was able to recover from leaf water potentials as low as -92 bar. The degree of stress experienced during the single drying cycle did not influence the maximum net photosynthetic rate attained during recovery, but the time taken to reach the maximum increased with the degree of stress experienced. During the first 24 h, the rate of recovery of net photosynthesis was mainly determined by the rate at which the water status improved. Leaves which experienced water potentials less than c. -40 bar had a slower rate of recovery of water potential than less stressed leaves. This was partially offset by higher rates of net photosynthesis. Furthermore, the relationship between leaf water potential and net photosynthesis recorded during the drying cycle was different from those measured during recovery. Thus different relationships must be used in models simulating behaviour during water stress and subsequent recovery. Stomatal resistance exerted greater control than intracellular resistance over net photosynthesis in the recovery phase, irrespective of the water potential before rewatering or whether plants were preconditioned to stress. Although abscisic acid concentration was positively related to leaf water potential and stomatal resistance during the drying cycle, the relationship between abscisic acid concentration and stomatal resistance during recovery was poor or absent. Sucrose and amino-acid nitrogen accumulated during stress and decreased during recovery. However, the level of non- structural carbohydrates or nitrogen compounds in the recovery phase did not appear to influence net photosynthetic rate or its components. In fact, the reverse appeared to occur: the rate of photosynthesis and growth seemed to determine the levels of these compounds.

Three tropical grasses, green panic (Panicum maximum var, trichoglume), spear grass (Heteropogon contortus) and buffel grass (Cenchrus ciliaris) and the tropical legume siratro (Macroptilium atropurpureum), were grown in plots in a semi-arid field environment. The water relations characteristics of leaves from plants subjected to a soil drying cycle were compared with those of unstressed leaves from plants in irrigated plots. Minimum water potentials attained in the stressed leaves were c. -44, - 38, - 33 and - 13 bar for the four species, respectively. The grass leaves adjusted osmotically to water stress, apparently through accumulation of solutes, so that there was a decrease in osmotic potential at full turgor (Ψπ100) of 5.5, 3.9 and 7.1 bar, and in water potential at zero turgor (Ψ0) of 8.6, 6.5 and 8.6 bar for green panic, spear grass and buffel respectively. Water stress appeared to increase slightly the proportion of bound water (B) and the bulk modulus of elasticity (ε) of the grass leaves, but it did not alter the relative water content at zero turgor (RWC0) or the ratio of turgid water content to dry weight of the tissue. The Ψπ100 and Ψ0 of stressed siratro leaves decreased by 2.5-4 bar and 3-5 bar respectively when subjected to soil drying cycles. These changes could be explained by the marked decrease in the ratio of turgid water content to dry weight of the leaf tissue rather than by accumulation of solutes. The values of RWC0 and ε for siratro leaves were not altered by stress but, in contrast to the grasses, B was apparently decreased although the data exhibited high variability. Adjustments in Ψπ100 and Ψ0 of stressed leaves of buffel grass and siratro were largely lost within 10 days of rewatering.

Different levels of stress were induced in P. maximum var. trichoglume by withholding water for various periods. Leaf elongation rate was measured during dark stress periods and after rewatering in both the light and the dark. Following rewatering, elongation rates of previously stressed plants exceeded those of controls for periods up to 33 h, during which time elongation rate was more related to previous levels of water stress than to current leaf water potential. In addition, there was a transient burst of elongation when plants were rewatered in the light. It is suggested that the stimulated rates result from expansion of cells which have accumulated during the stress because cell division is less sensitive to water stress than is cell expansion. Despite the stimulated rates of elongation after rewatering, the recovery was incomplete such that the final lengths of stressed leaves were less than those of unstressed plants.

The responses of carbon dioxide exchange and leaf elongation of potted P. maximum var. trichoglume plants to water deficits were investigated in controlled environments and outdoors during drying cycles down to -92 bars leaf water potential, The sensitivities of net photosynthesis and leaf elongation to water deficits were similar. The leaf water potentials at which net photosynthesis and elongation ceased (c. -12 bars), and stomatal resistance increased substantially (- 6 bars), were relatively unaffected by nitrogen supply, environmental conditions during growth, and whether plants had previously experienced stress. However, these factors influenced the rate of net photosynthesis, at high leaf water potentials by affecting stomatal resistance and at moderate water potentials by affecting both stomatal and intracellular resistances. Stomata1 resistance was more sensitive than intracellular resistance to water deficits. Dark respiration rate decreased with leaf water potential, and was higher in plants receiving additional nitrogen. At moderate leaf water potentials (-7 to -9 bars), net photosynthesis of this C4 grass exhibited light saturation and rates similar to C3 plants. We suggest that the difference in behaviour of controlled-environment-grown and field-grown plants to water deficits observed with some species is unlikely to be due to differences in the aerial environment, but may result from differences in the rate at which stress develops. The ecological significance and evolution of the C4 syndrome are discussed briefly.

The effects of recurring cycles of short-term water stress on the water relations and growth of P. maximum var. trichoglume in pots of soil were investigated under controlled conditions. As soil water content decreased there was an increase in the resistance to water movement in the soil-plant system. Leaf stomatal resistance increased and concomitantly transpiration rate decreased when soil water content fell below 37 % (soil water potential of - 1 .0 bars) and leaf water potentials were less than - 6 bars. The leaf water potential at wilting (- 8 to - 10 bars) and the relation between leaf water potential and relative water content changed with leaf position on the tiller. The death of early-formed leaves on the plants was accelerated by water stress but, in contrast, the later-formed leaves died more rapidly in the control (unstressed) treatment so that finally the control plants had a higher proportion of dead leaves. Plant growth was reduced at soil water contents above the permanent wilting point. Reduction in net assimilation rate was the main determinant of lower relative growth rate of stressed plants over the initial cycles of stress but subsequently, as leaf area expansion was reduced, leaf area ratio also had a significant influence. Water stress influenced growth directly, and also indirectly via its effect on plant development (ontogeny). Two techniques were used to separate the direct from the indirect effects on relative growth rate Some published data which suggest a stimulation of growth rate after the relief of stress are re-interpreted and the effect is shown to be due mainly to differences in ontogeny between stressed and control treatments

A previous study of one grass and one legume at early vegetative stages has been extended to include a further nine species of each. It confirms the superior relative growth rates (RW) of grasses, arising from very high net assimilation rates (EA), in turn dependent on high photosynthetic rates (P). Leaf area ratios (FA) varied over a similar range in both groups. RW values of grasses in the second week after sowing varied from 0.41 to 0.55 gg-1 day-1; for legumes, they were 0.31-0.36. Corresponding EA values were 1.5-1.9 g dm-2 wk-1 in grasses, and 1.1-1.8 in legumes. Within both grasses and legumes, comparisons between species can be made with less certainty. In grasses, variation in FA combined with generally uniform EA values resulted in variable RW values. Both FA and EA varied in the legumes but were usually compensatory, leading to uniformity in RW. Watson and Hayashi's method of separating EA into photosynthetic and respiratory components was used. Differences both between and within grasses and legumes were shown, but there are probably important errors in this technique, particularly where there are discrepancies between light and dark respiration.

Hamil grass and Siratro were grown as single plants in pots in controlled environment cabinets at a temperature favouring high growth rates and with adequate water and mineral nutrients. Leaf areas and dry weights of plant parts were recorded in two experiments, to which some of the plants were common. In the first experiment, weekly samples until 4 weeks from sowing provided data for the calculation of relative growth rates, net assimilation rates, and leaf area ratios. The relative growth rate of Hamil grass was almost twice that of Siratro and appears to be the highest yet recorded for any plant. The higher growth rate for the grass resulted from a high net assimilation rate, although the leaf area ratio was lower. The second experiment used Watson and Hayashi's method, in which plants are kept in darkness for a varying number of days to separate net assimilation rate into photosynthetic and respiratory components. Data were collected at 2 and 4 weeks from sowing. The high net assimilation rate of the grass was shown to result from a higher photosynthetic rate, despite a much greater respiration rate than in the legume. The technique used for estimating photosynthesis and respiration is discussed in relation to some of the assumptions that it makes.