As superficial structures, non-glandular trichomes, protect plant organs against multiple biotic and abiotic stresses. The protective and defensive roles of these epidermal appendages are crucial to developing organs and can be attributed to the excellent combination of suitable structural traits and chemical reinforcement in the form of phenolic compounds, primarily flavonoids. Both the formation of trichomes and the accumulation of phenolics are interrelated at the molecular level. During the early stages of development, non-glandular trichomes show strong morphological similarities to glandular ones such as the balloon-like apical cells with numerous phenolics. At later developmental stages, and during secondary wall thickening, phenolics are transferred to the cell walls of the trichomes. Due to the diffuse deposition of phenolics in the cell walls, trichomes provide protection against UV-B radiation by behaving as optical filters, screening out wavelengths that could damage sensitive tissues. Protection from strong visible radiation is also afforded by increased surface light reflectance. Moreover, the mixtures of trichome phenolics represent a superficial chemical barrier that provides protection against biotic stress factors such as herbivores and pathogens. Although the cells of some trichomes die at maturity, they can modulate their quantitative and qualitative characteristics during development, depending on the prevailing conditions of the external biotic or abiotic environment. In fact, the structure and chemical constituents of trichomes may change due to the particular light regime, herbivore damage, wounding, water stress, salinity and the presence of heavy metals. Hence, trichomes represent dynamic protective structures that may greatly affect the outcome of many plant–environment interactions.

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 pigweed (Amaranthus hybridus) 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 nonatmospheric 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 CO 2 . Similar results were also observed under drought stress in Dianthus chinensis, Pelargonium peltatum, and Portulacaria afra. Moreover, in A. hybridus, 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 nonatmospheric 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.

Carbon-calcium inclusions (CCaI) either as calcium oxalate crystals (CaOx) or amorphous calcium carbonate cystoliths are spread among most photosynthetic organisms. They represent dynamic structures with a significant construction cost and their appearance during evolution indicates an ancient origin. Both types of inclusions share some similar functional characteristics providing adaptive advantages such as the regulation of Ca levels, and the release of CO 2 and water molecules upon decomposition. The latter seems to be essential under drought conditions and explains the intense occurrence of these structures in plants thriving in dry climates. It seems, however, that for plants CaOx may represent a more prevalent storage system compared with CaCO 3 due to the multifunctionality of oxalate. This compound participates in a number of important soil biogeochemical processes, creates endosymbiosis with beneficial bacteria and provides tolerance against a combination of abiotic (nutrient deprivation, metal toxicity) and biotic (pathogens, herbivores) stress factors. We suggest a re-evaluation of the roles of these fascinating plant structures under a new and holistic approach that could enhance our understanding of carbon sequestration at the whole plant level and provide future perspectives.

Summary Mesophyll conductance (gm), the diffusion of CO2 from substomatal cavities to the carboxylation sites in the chloroplasts, is a highly complex trait driving photosynthesis (net CO2 assimilation, AN). However, little is known concerning the mechanisms by which it is dynamically regulated. The apoplast is considered as a ‘key information bridge’ between the environment and cells. Interestingly, most of the environmental constraints affecting gm also cause apoplastic responses, cell wall (CW) alterations and metabolic rearrangements. Since CW thickness is a key determinant of gm, we hypothesize that other changes in this cellular compartiment should also influence gm. We study the relationship between the antioxidant apoplastic system and CW metabolism and the gm responses in tobacco plants (Nicotiana sylvestris L.) under two abiotic stresses (drought and salinity), combining in vivo gas‐exchange measurements with analyses of antioxidant activities, CW composition and primary metabolism. Stress treatments imposed substantial reductions in AN (58–54%) and gm (59%), accompanied by a strong antioxidant enzymatic response at the apoplastic and symplastic levels. Interestingly, apoplastic but not symplastic peroxidases were positively related to gm. Leaf anatomy remained mostly stable; however, the stress treatments significantly affected the CW composition, specifically pectins, which showed significant relationships with AN and gm. The treatments additionally promoted a differential primary metabolic response, and specific CW‐related metabolites including galactose, glucosamine and hydroxycinnamate showed exclusive relationships with gm independent of the stress. These results suggest that gm responses can be attributed to specific changes in the apoplastic antioxidant system and CW metabolism, opening up more possibilities for improving photosynthesis using breeding/biotechnological strategies.

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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.

Water deprivation affects photosynthesis, leaf anatomy and cell wall composition. Although the former effects have been widely studied, little is known regarding those changes in cell wall major (cellulose, hemicelluloses, pectin and lignin) and minor (cell wall bound phenolics) compounds in plants acclimated to short- and long-term water deprivation and during recovery. Particularly, how these cell wall changes impact anatomy and/or photosynthesis, specifically mesophyll conductance to CO2 diffusion (gm), has been scarcely studied. To induce changes in photosynthesis, cell wall composition and anatomy, Helianthus annuus plants were studied under five conditions: (i) control (i.e., without stress) (“CL”), (ii) long-term water deficit stress (“LT”), (iii) long-term water deficit stress with recovery (“LT-Rec”), (iv) short-term water deficit stress (“ST”) and (v) short-term water deficit stress with recovery (“ST-Rec”), resulting in a wide photosynthetic range (from 3.80 ± 1.05 to 24.53 ± 0.42 μmol CO2 m -2 s -1). Short- and long-term water deprivation and recovery induced distinctive responses of the examined traits, evidencing a cell wall dynamic turnover during plants acclimation to each condition. Particularly, we demonstrated for the first time how gm correlated negatively with lignin and cell wall bound phenolics and how (Cellulose+Hemicelloses)/Pectin ratio was linked to cell wall thickness (Tcw) variations.

This study reveals a negative relationship between leaf phenolic compounds and photosynthetic Amax among different plant species. This indicates a functional integration among carbon gain and the concentration of leaf phenolic compounds that reflects the trade-off between growth and defence/protection demands.