The literature and our present examinations indicate that the intra-leaf light absorption profile is in most cases steeper than the photosynthetic capacity profile. In strong white light, therefore, the quantum yield of photosynthesis would be lower in the upper chloroplasts, located near the illuminated surface, than that in the lower chloroplasts. Because green light can penetrate further into the leaf than red or blue light, in strong white light, any additional green light absorbed by the lower chloroplasts would increase leaf photosynthesis to a greater extent than would additional red or blue light. Based on the assessment of effects of the additional monochromatic light on leaf photosynthesis, we developed the differential quantum yield method that quantifies efficiency of any monochromatic light in white light. Application of this method to sunflower leaves clearly showed that, in moderate to strong white light, green light drove photosynthesis more effectively than red light. The green leaf should have a considerable volume of chloroplasts to accommodate the inefficient carboxylation enzyme, Rubisco, and deliver appropriate light to all the chloroplasts. By using chlorophylls that absorb green light weakly, modifying mesophyll structure and adjusting the Rubisco/chlorophyll ratio, the leaf appears to satisfy two somewhat conflicting requirements: to increase the absorptance of photosynthetically active radiation, and to drive photosynthesis efficiently in all the chloroplasts. We also discuss some serious problems that are caused by neglecting these intra-leaf profiles when estimating whole leaf electron transport rates and assessing photoinhibition by fluorescence techniques.
Dynamic acclimation of the photosynthetic apparatus in response to environmental cues, particularly light quantity and quality, is a widely-observed and important phenomenon which contributes to the tolerance of plants against stress and helps to maintain, as far as possible, optimal photosynthetic efficiency and resource utilization. This mini-review represents a scrutiny of a number of possible photoreceptors (including the two photosystems acting as light sensors) and signal transducers that may be involved in producing acclimation responses. We suggest that regulation by signal transduction may be effected at each of several possible points, and that there are multiple regulatory mechanisms for photosynthetic acclimation.
We propose a simplified alternative method for quantifying the partitioning of excitation energy between photochemistry, fluorescence and thermal dissipation. This alternative technique uses existing well-defined quantum efficiencies such as Phi(PS II), leaving no 'excess' efficiency unaccounted for, effectively separates regulated and constitutive thermal dissipation processes, does not require the use of F(o) and F'(o) measurements and gives very similar results to the method proposed by Kramer et al. [(2004) Photosynth Res 79: 209-218]. We demonstrate the use of the technique using chlorophyll fluorescence measurements in grapevine leaves and observe a high dependence on thermal dissipation processes (up to 75%) at both high light and low temperature.
In 27 C 4 grasses grown under adequate or deficient nitrogen (N) supplies, N-use efficiency at the photosynthetic (assimilation rate per unit leaf N) and whole-plant (dry mass per total leaf N) level was greater in NADP-malic enzyme (ME) than NAD-ME species. This was due to lower N content in NADP-ME than NAD-ME leaves because neither assimilation rates nor plant dry mass differed significantly between the two C 4 subtypes. Relative to NAD-ME, NADP-ME leaves had greater in vivo (assimilation rate per Rubisco catalytic sites) and in vitro Rubisco turnover rates (k cat ; 3.8 versus 5.7 s 21 at 25°C). The two parameters were linearly related. In 2 NAD-ME (Panicum miliaceum and Panicum coloratum) and 2 NADP-ME (Sorghum bicolor and Cenchrus ciliaris) grasses, 30% of leaf N was allocated to thylakoids and 5% to 9% to amino acids and nitrate. Soluble protein represented a smaller fraction of leaf N in NADP-ME (41%) than in NAD-ME (53%) leaves, of which Rubisco accounted for one-seventh. Soluble protein averaged 7 and 10 g (mmol chlorophyll) 21 in NADP-ME and NAD-ME leaves, respectively. The majority (65%) of leaf N and chlorophyll was found in the mesophyll of NADP-ME and bundle sheath of NAD-ME leaves. The mesophyll-bundle sheath distribution of functional thylakoid complexes (photosystems I and II and cytochrome f ) varied among species, with a tendency to be mostly located in the mesophyll. In conclusion, superior N-use efficiency of NADP-ME relative to NAD-ME grasses was achieved with less leaf N, soluble protein, and Rubisco having a faster k cat .C 4 photosynthesis involves the close collaboration of two photosynthetic cell types, the mesophyll (M) and bundle sheath (BS). A key characteristic of the C 4 syndrome is the operation of a CO 2 concentrating mechanism, which serves to raise the CO 2 concentration in the BS around Rubisco to levels high enough to suppress photorespiration and almost saturate photosynthesis in air (Hatch, 1987). This explains the commonly observed high photosynthetic rates of C 4 relative to C 3 leaves, when comparisons are made under high light and temperature. C 4 plants also have greater photosynthetic rates and accumulate more biomass than C 3 plants for less leaf nitrogen (N) and Rubisco (Bolton and Brown, 1978;Brown, 1978;Schmitt and Edwards, 1981;Ghannoum et al., 1997;Ghannoum and Conroy, 1998;Makino et al., 2003). The C 4 photosynthetic pathway is divided into three biochemical subtypes following the major C 4 acid decarboxylation enzyme (NAD-malic enzyme [ME], NADP-ME, and phosphoenolpyruvate carboxykinase; Hatch, 1987). C 4 grasses with different biochemical subtypes have characteristic leaf anatomy (Hattersley, 1992) and different geographic distribution according to rainfall, such as seen in Australia (Hattersley, 1992) and South Africa (Ellis et al., 1980). With increasing rainfall, NADP-ME grasses increase in abundance, whereas NAD-ME grasses become less abundant. The aforementioned observations triggered our interest in the comparative physiology of the C 4 subtypes, espec...
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