The photosynthetic capacity of leaves is related to the nitrogen content primarily bacause the proteins of the Calvin cycle and thylakoids represent the majority of leaf nitrogen. To a first approximation, thylakoid nitrogen is proportional to the chlorophyll content (50 mol thylakoid N mol-1 Chl). Within species there are strong linear relationships between nitrogen and both RuBP carboxylase and chlorophyll. With increasing nitrogen per unit leaf area, the proportion of total leaf nitrogen in the thylakoids remains the same while the proportion in soluble protein increases. In many species, growth under lower irradiance greatly increases the partitioning of nitrogen into chlorophyll and the thylakoids, while the electron transport capacity per unit of chlorophyll declines. If growth irradiance influences the relationship between photosynthetic capacity and nitrogen content, predicting nitrogen distribution between leaves in a canopy becomes more complicated. When both photosynthetic capacity and leaf nitrogen content are expressed on the basis of leaf area, considerable variation in the photosynthetic capacity for a given leaf nitrogen content is found between species. The variation reflects different strategies of nitrogen partitioning, the electron transport capacity per unit of chlorophyll and the specific activity of RuBP carboxylase. Survival in certain environments clearly does not require maximising photosynthetic capacity for a given leaf nitrogen content. Species that flourish in the shade partition relatively more nitrogen into the thylakoids, although this is associated with lower photosynthetic capacity per unit of nitrogen.

Photosynthesis and nitrogen relationships in leaves of C3 plants.

Publisher Summary This chapter focuses on evolutionary significance of phenotypic plasticity in plants. The expression of an individual genotype is modified by its environment. The amount by which it can be modified is termed its plasticity. This plasticity can be either morphological or physiological; these are interrelated. The plasticity of a character is related to the general pattern of its development, and apart from this, that plasticity is a general property of the whole genotype. Plasticity of a character appears to be specific for that character, specific in relation to particular environmental influences, specific in direction, under genetic control not necessarily related to heterozygosity, and able to be radically altered by selection. Because plants are static organisms, plasticity is of marked adaptive value in a great number of situations. Examples of all these situations in plant species are discussed. They indicate that adaptation by plasticity is a widespread and important phenomenon in plants and has evolved differently in different species. The mechanisms involved in plasticity are varied. At one extreme, the character shows a continuous range of modification dependent on the intensity of the environmental stimulus. At the other, the character shows only two discrete modifications. The stimulus causing these modifications may be direct or indirect. The mechanisms found can be related to the particular environmental situation involved.

Evolutionary Significance of Phenotypic Plasticity in Plants

INTRODUCTION 492 ECOLOGICAL ASPECTS OF PHOTOSYNTHETIC TEMPERATURE ADAPTATION 493 Photosynthetic Temperature Dependence in Thermally Contrasting Climates ........ 493 Photosynthetic Temperature Acclimation 497 Seasonal acclimation in natural habitats ...... ....... 497 Studies in controlled environments 499 THE MECHANISTIC BASIS FOR PHOTOSYNTHETIC RESPONSE AND ADAPTATION TO TEMPERATURE 504 Reversible Temperature Respon.ses 505 Stomata! effec� o� the . temJH!.rature response 0/ photo.rynthesis ......... ....... 505 Interacttons with /lght mtenslty 507 C, photo.rynthesis (lM photorespiration 507 C, photo.rynthesis 515 Comparison 0/ plants from contrasting thermol regimes ...... ...... 517 ["eversible Temperature Respon.ses 519 Low temperature sensitMty ....... 519 High temperature sensitivity 524 Adoptive responses in the heat stability 0/ the photosynthetic apparatus 530 CONCLUDING REMARKS 532

Photosynthetic Response and Adaptation to Temperature in Higher Plants

▪ Abstract The primary effect of the response of plants to rising atmospheric CO2 (Ca) is to increase resource use efficiency. Elevated Ca reduces stomatal conductance and transpiration and improves water use efficiency, and at the same time it stimulates higher rates of photosynthesis and increases light-use efficiency. Acclimation of photosynthesis during long-term exposure to elevated Ca reduces key enzymes of the photosynthetic carbon reduction cycle, and this increases nutrient use efficiency. Improved soil–water balance, increased carbon uptake in the shade, greater carbon to nitrogen ratio, and reduced nutrient quality for insect and animal grazers are all possibilities that have been observed in field studies of the effects of elevated Ca. These effects have major consequences for agriculture and native ecosystems in a world of rising atmospheric Ca and climate change.

More Efficient Plants : a consequence of rising atmospheric CO2

Whole-plant energy capture depends not only on the photosynthetic response of individual leaves, but also on their integration into an effective canopy, and on the costs of producing and maintaining their photosynthetic capacity. This paper explores adaptation to irradiance level in this context, focusing on traits whose significance would be elusive if considered in terms of their impact at the leaf level alone. I review traditional approaches used to demonstrate or suggest adaptation to irradiance level, and outline three energetic tradeoffs likely to shape such adaptation, involving the economics of gas exchange, support, and biotic interactions. Recent models using these tradeoffs to account for trends in leaf nitrogen content, stornatal conductance, phyllotaxis, and defensive allocations in sun v. shade are evaluated. A re-evaluation of the classic study of acclimation of the photosynthetic light response in Atriplex, crucial to interpreting adaptation to irradiance in many traits, shows that it does not completely support the central dogma of adaptation to sun v. shade unless the results are analysed in terms of whole-plant energy capture. Calculations for Liriodendron show that the traditional light compensation point has little meaning for net carbon gain, and that the effective compensation point is profoundly influenced by the costs of night leaf respiration, leaf construction, and the construction of associated support and root tissue. The costs of support tissue are especially important, raising the effective compensation point by 140 pmol m- s - ' in trees 1 m tall, and by nearly 1350 pmol m - s - ' in trees 30 m tall. Effective compensation points give maximum tree heights as a function of irradiance, and shade tolerance as a function of tree height; calculations of maximum permissible height in Liriodendron correspond roughly with the height of the tallest known individual. Finally, new models for the evolution of canopy width/height ratio in response to irradiance and coverage within a tree stratum, and for the evolution of mottled leaves as a defensive measure in understory herbs, are outlined.

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Adaptation to Sun and Shade: a Whole-Plant Perspective

Adaptability of the Photosynthetic Apparatus to Light Intensity in Ecotypes from Exposed and Shaded Habitats

Photosynthetic adaptation to light intensity has been studied in clones of populations from shaded and exposed habitats of Rumex acetosa and Geum rivale. Clones of the shade species Lamium galeobdolon and the sun species Plantago lanceolata were also included for comparison. The plants were grown under controlled conditions at a high and a low light intensity. The capacity of photosynthetic carbon dioxide uptake at low as well as at saturating light intensities was determined on single attached leaves.



As was previously demonstrated in Solidago virgaurea, clones of populations native to shaded and to exposed environments show differences in the photosynthetic response to light intensity during growth. The data provide evidence that populations of the same species native to habitats with contrasting light intensities differ in their photosynthetic properties in an adaptive manner Ln a similar mode as sun and shade species.

Photosynthetic Adaptation to Light Intensity in Plants Native to Shaded and Exposed Habitats

Plants of Solidago virgaurea L. from exposed and shaded habitats differ with respect to the response of the photosynthetic apparatus to the level of irradiance during growth. An analysis was carried out on leaf characteristies which might be responsible for the differences established in the rates of Hght-saturated CO2 uptake. The clones were grown in controlled environment chambers at high and low levels of irradiance. Light-saturated rates of photosynthesis and transpiration were measured at natural and lower ambient CO2 concentrations.

A low temperature dependence of light-saturated CO2 uptake at natural CO2 concentrations, and a strong response to changes in stomatal width, suggested that the rate of CO2 transfer from ambient air towards reaetion sites in chloroplasts was mainly limiting the pholosynthetic rate. Resistances to transfer of CO2 for different parts of the pathway were calculated. There was a weak but significant correlation between stomatal conductance and the product stomatal frequency ± pore length. Mesopbyll conductance and dry weight per unit area were highly correlated in leaves not damaged by high irradiance. This suggests that mesophyll conductance increases with increasing cross sectional area (per unit leaf area) of the pathways of CO2 transfer in the mesophyll from cell surfaces to reaction sites.

The higher light-saturated photosynthesis in clones from exposed habitats when grown at high irradiance than when grown at low irradiance was attributable mainly to a lower mesophyll resistance. In shade clones the effect upon CO2 uptake of the increase in leaf thickness when grown at high irradiance was counteracted by the associated inactivation of the photosynthetic apparatus. The difference in CO2 uptake present between clones from exposed and shaded habitats when preconditioned to high irradiance resulted from differences in both mesophyll and stomatal resistances. A few hybrid clones of an F1-population from a cross between a clone from an exposed habitat and a clone from a shaded habitat reacted, on the whole, in the same way as the exposed habitat parent. When grown at high irradiance, the hybrid clones showed higher photosynthetic rates than either parent; this was largely attributable to the unusually low stomatal resistance of the hybrid leaves.

Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Efflux of carbon dioxide in light and darkness was measured at low ambient CO2 concentrations in leaves of Rumex acetosa. Light carbon dioxide production (photo-respiration) was found to depend on irradiance and to differ from dark production as to the response to temperature and ambient concentrations of O2 and CO2. These observations support previously made suggestions that photorespiration follows a different metabolic pathway to dark respiration.

Paul Holmgren

Papers

Adaptability of the Photosynthetic Apparatus to Light Intensity in Ecotypes from Exposed and Shaded Habitats

Photosynthetic Adaptation to Light Intensity in Plants Native to Shaded and Exposed Habitats

Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Carbon Dioxide Efflux from Leaves in Light and Darkness

Polyphenols and Polyphenol Oxidases in Leaves of Solidago virgaurea