Calcium oxalate crystals are widespread among animals and plants. In land plants, crystals often reach high amounts - up to 80% of dry biomass. They are formed within specific cells, and their accumulation constitutes a normal activity rather than a pathological symptom, as occurs in animals. Despite their ubiquity, our knowledge on the formation and the possible role(s) of these crystals remains limited. We show that the mesophyll crystals of Amaranthus hybridus (pigweed) exhibit diurnal volume changes with a gradual decrease during daytime and a total recovery during the night. Moreover, stable carbon isotope composition indicated that crystals are of non-atmospheric origin. Stomatal closure (under drought conditions or exogenous application of abscisic acid) was accompanied by crystal decomposition and by increased activity of oxalate oxidase that converts oxalate into CO2. Similar results were also observed under drought stress in Dianthus chinensis, Pelargonium peltatum and Portulacaria afra. Moreover, in pigweed, despite closed stomata, the leaf metabolic profiles combined with chlorophyll fluorescence measurements indicated active photosynthetic metabolism. In combination, calcium oxalate crystals in leaves can act as a biochemical reservoir that collects non-atmospheric carbon, mainly during the night. During the day, crystal degradation provides subsidiary carbon for photosynthetic assimilation, especially under drought conditions. This new photosynthetic path, with the suggested name "alarm photosynthesis", seems to provide a number of adaptive advantages such as water economy, limitation of carbon losses to the atmosphere and a lower risk of photoinhibition, roles that justify its vast presence in plants.
• Background and Aims Depending on cultivar, surfaces of young leaves of Vitis vinifera may be glabrous-green (‘Soultanina’) or transiently have anthocyanins (‘Siriki’) or pubescence (‘Athiri’). A test is made of the hypothesis that anthocyanins and pubescence act as light screens affording a photoprotective advantage to the corresponding leaves, and an assessment is made of the magnitude of their effect. • Methods Measurements were made on young leaves of the three cultivars in spring under field conditions. Photosynthetic gas-exchange and in vivo chlorophyll fluorescence were measured. Photosynthetic and photoprotective pigments were analysed by HPLC. • Key Results Compared with glabrous-green leaves, both anthocyanic and pubescent leaves had greater dark-adapted PSII photochemical efficiency and net photosynthesis. In leaves possessing either anthocyanins or pubescence, the ratio of xanthophyll cycle components to total chlorophyll, and mid-day de-epoxidation state of the xanthophyll cycle were considerably smaller, than in glabrous-green leaves. These differences were more evident in pubescent leaves, probably indicating that trichomes were more effective in decreasing light stress than anthocyanins in the epidermis. • Conclusions Light screens, especially in the form of pubescence, decrease the risk of photoinhibition whilst allowing leaves to maintain a smaller content of xanthophyll cycle components and depend less on xanthophyll cycle energy dissipation. This combination of photoprotective features, i.e. decreased photon flux to the photosynthetic apparatus and lower xanthophyll cycle utilization rates may be particularly advantageous under stressful conditions.
Leaves have evolved to effectively harvest light, and, in parallel, to balance photosynthetic CO2 assimilation with water losses. At times, leaves must operate under light limiting conditions while at other instances (temporally distant or even within seconds), the same leaves must modulate light capture to avoid photoinhibition and achieve a uniform internal light gradient. The light-harvesting capacity and the photosynthetic performance of a given leaf are both determined by the organization and the properties of its structural elements, with some of these having evolved as adaptations to stressful environments. In this respect, the present review focuses on the optical roles of particular leaf structural elements (the light capture module) while integrating their involvement in other important functional modules. Superficial leaf tissues (epidermis including cuticle) and structures (epidermal appendages such as trichomes) play a crucial role against light interception. The epidermis, together with the cuticle, behaves as a reflector, as a selective UV filter and, in some cases, each epidermal cell acts as a lens focusing light to the interior. Non glandular trichomes reflect a considerable part of the solar radiation and absorb mainly in the UV spectral band. Mesophyll photosynthetic tissues and biominerals are involved in the efficient propagation of light within the mesophyll. Bundle sheath extensions and sclereids transfer light to internal layers of the mesophyll, particularly important in thick and compact leaves or in leaves with a flutter habit. All of the aforementioned structural elements have been typically optimized during evolution for multiple functions, thus offering adaptive advantages in challenging environments. Hence, each particular leaf design incorporates suitable optical traits advantageously and cost-effectively with the other fundamental functions of the leaf.
The surface of peach (Prunus persica ‘Calrico’) is covered by a dense indumentum, which may serve various protective purposes. With the aim of relating structure to function, the chemical composition, morphology, and hydrophobicity of the peach skin was assessed as a model for a pubescent plant surface. Distinct physicochemical features were observed for trichomes versus isolated cuticles. Peach cuticles were composed of 53% cutan, 27% waxes, 23% cutin, and 1% hydroxycinnamic acid derivatives (mainly ferulic and p-coumaric acids). Trichomes were covered by a thin cuticular layer containing 15% waxes and 19% cutin and were filled by polysaccharide material (63%) containing hydroxycinnamic acid derivatives and flavonoids. The surface free energy, polarity, and work of adhesion of intact and shaved peach surfaces were calculated from contact angle measurements of water, glycerol, and diiodomethane. The removal of the trichomes from the surface increased polarity from 3.8% (intact surface) to 23.6% and decreased the total surface free energy chiefly due to a decrease on its nonpolar component. The extraction of waxes and the removal of trichomes led to higher fruit dehydration rates. However, trichomes were found to have a higher water sorption capacity as compared with isolated cuticles. The results show that the peach surface is composed of two different materials that establish a polarity gradient: the trichome network, which has a higher surface free energy and a higher dispersive component, and the cuticle underneath, which has a lower surface free energy and higher surface polarity. The significance of the data concerning water-plant surface interactions is discussed within a physiological context.