The Orchidaceae is a diverse and wide spread family of flowering plants that are of great value in ornamental, medical, conservation, and evolutionary research. The broad diversity in morphology, growth form, life history, and habitat mean that the members of Orchidaceae exhibit various physiological properties. Epiphytic orchids are often characterized by succulent leaves with thick cell walls, cuticles, and sunken stomata, whereas terrestrial orchids possess rhizomes, corms, or tubers. Most orchids have a long juvenile period, slow growth rate, and low photosynthetic capacity. This reduced photosynthetic potential can be largely explained by CO2 diffusional conductance and leaf internal structure. The amount of light required for plant survival depends upon nutritional mode, growth form, and habitat. Most orchids can adapt to their light environments through morphological and physiological adjustments but are sensitive to sudden changes in irradiance. Orchids that originate from warm regions are susceptible to chilling temperatures, whereas alpine members are vulnerable to high temperatures. For epiphytic orchids, rapid water uptake by the velamen radicum, water storage in their pseudobulbs and leaves, slow water loss, and Crassulacean Acid Metabolism contribute to plant-water balance and tolerance to drought stress. The presence of the velamen radicum and mycorrhizal fungi may compensate for the lack of root hairs, helping with quick absorbance of nutrients from the atmosphere. Under cultivation conditions, the form and concentration of nitrogen affect orchid growth and flowering. However, the limitations of nitrogen and phosphorous on epiphytic orchids in the wild, which require these plants to depend on mycorrhizal fungi for nutrients throughout the entire life cycle, are not clearly understood. Because they lack endosperm, seed germination depends upon obtaining nutrients via mycorrhizal fungi. Adult plants of some autotrophic orchids also gain carbon, nitrogen, phosphorus, and other elements from their mycorrhizal partners. Future studies should examine the mechanisms that determine slow growth and flower induction, the physiological causes of variations in flowering behavior and floral lifespan, the effects of nutrients and atmospheric-nitrogen deposition, and practical applications of mycorrhizal fungi in orchid cultivation.

In higher plants, the generation of proton gradient across the thylakoid membrane (pH) through cyclic electron flow (CEF) has mainly two functions: 1) to generate ATP and balance the ATP/NADPH energy budget, and 2) to protect photosystems I and II against photoinhibition. The intensity of light under which plants are grown alters both CEF activity and the ATP/NADPH demand for primary metabolic processes. However, it is unclear how the role of CEF is affected by the level of irradiance that is applied during the growth and measurement periods. We studied the role of CEF at different light intensities in leaves from sun- and shade-grown plants. At 849 μmol photons m-2 s-1, both types of leaves had nearly the same degree of CEF activation. Modeling of the ATP/NADPH demand revealed that, at this light intensity, the contribution of CEF toward supplying ATP was much higher in the sun leaves. Meanwhile, the shade leaves showed higher levels of non-photochemical quenching and the P700 oxidation ratio. Therefore, at 849 μmol photons m-2 s-1, CEF mainly helped in the synthesis of ATP in the sun leaves, but functioned in photoprotection for the shade leaves. When the light intensity increased to 1976 μmol photons m-2 s-1, CEF activation was greatly enhanced in the sun leaves, but its contribution to supplying ATP changed slightly. These results indicate that the main role of CEF is altered flexibly in response to light intensity. In particular, CEF mainly contributes to balancing the ATP/NADPH energy budget under sub-saturating light intensities. When exposed to saturating light intensities, CEF mainly protects photosynthetic apparatus against photoinhibition.

Plants usually experience dynamic fluctuations of light intensities under natural conditions. However, the responses of mesophyll conductance, CO2 assimilation, and photorespiration to light fluctuation are not well understood. To address this question, we measured photosynthetic parameters of gas exchange and chlorophyll fluorescence in tobacco leaves at 2-min intervals while irradiance levels alternated between 100 and 1200 μmol photons m-2 s-1. Compared with leaves exposed to a constant light of 1200 μmol photons m-2 s-1, both stomatal and mesophyll conductances were significantly restricted in leaves treated with fluctuating light condition. Meanwhile, CO2 assimilation rate and electron flow devoted to RuBP carboxylation at 1200 μmol photons m-2 s-1 under fluctuating light were limited by the low chloroplast CO2 concentration. Analysis based on the C3 photosynthesis model indicated that, at 1200 μmol photons m-2 s-1 under fluctuating light, the CO2 assimilation rate was limited by RuBP carboxylation. Electron flow devoted to RuBP oxygenation at 1200 μmol photons m-2 s-1 under fluctuating light remained at nearly the maximum level throughout the experimental period. We conclude that fluctuating light restricts CO2 assimilation by decreasing both stomatal and mesophyll conductances. Under such conditions, photorespiration plays an important role in the regulation of photosynthetic electron flow.

Drought and heat stresses usually occur concomitantly in nature, with increasing frequency and intensity of both stresses expected due to climate change. The synergistic agricultural impacts of these compound climate extremes are much greater than those of the individual stresses. However, the mechanisms by which drought and heat stresses separately and concomitantly affect dynamic photosynthesis have not been thoroughly assessed. To elucidate this, we used tomato (Solanum lycopersicum) seedlings to measure dynamic photosynthesis under individual and compound stresses of drought and heat. Individual drought and heat stresses limited dynamic photosynthesis at the stages of diffusional conductance to CO₂ and biochemistry, respectively. However, the primary limiting factor for photosynthesis shifted to mesophyll conductance under the compound stresses. Compared with the control, photosynthetic carbon gain in fluctuating light decreased by 38%, 73%, and 114% under the individual drought, heat, and compound stresses, respectively. Therefore, compound stresses caused a greater reduction in photosynthetic carbon gain in fluctuating light conditions than individual stress. These findings highlight the importance of mitigating the effects of compound climate extremes on crop productivity by targeting mesophyll conductance and improving dynamic photosynthesis.

In higher plants, moderate photoinhibition of photosystem II (PSII) leads to a stimulation of cyclic electron flow (CEF) at low light, which is accompanied by an increase in the P700 oxidation ratio. However, the specific role of CEF stimulation at low light is not well known. Furthermore, the mechanism underlying this increase in P700 oxidation ratio at low light is unclear. To address these questions, intact leaves of the shade-adapted plant Panax notoginseng were treated at 2258 μmol photons m^-2 s^-1 for 30 min to induce PSII photoinhibition. Before and after this high-light treatment, PSI and PSII activity, the energy quenching in PSII, the redox state of PSI and proton motive force (pmf) at a low light of 54 μmol photons m^-2 s^-1 were determined at the steady state. After high-light treatment, electron flow through PSII (ETRII) significantly decreased but CEF was remarkably stimulated. The P700 oxidation ratio significantly increased but non-photochemical quenching changed negligibly. Concomitantly, the total pmf decreased significantly and the proton gradient (ΔpH) across the thylakoid membrane remained stable. Furthermore, the P700 oxidation ratio was negatively correlated with the value of ETRII. These results suggest that upon PSII photoinhibition, CEF is stimulated to increase the ATP synthesis, facilitating the rapid repair of photodamaged PSII. The increase in P700 oxidation ratio at low light cannot be explained by the change in pmf, but is primarily controlled by electron transfer from PSII.

Epiphytes that grow in the canopies of tropical and subtropical forests experience different water regimes when compared with terrestrial plants. However, the differences in adaptive strategies between epiphytic and terrestrial plants with respect to plant water relations remain poorly understood. To understand how water-related traits contrast between epiphytic and terrestrial growth forms within the Cymbidium (Orchidaceae), we assessed leaf anatomy, hydraulics, and physiology of seven terrestrial and 13 epiphytic species using a common garden experiment. Compared with terrestrial species, epiphytic species had higher values for leaf mass per unit area (LMA), leaf thickness (LT), epidermal thickness, saturated water content (SWC) and the time required to dry saturated leaves to 70% relative water content (T70). However, vein density (Dvein), stomatal density (SD), and photosynthetic capacity (Amax) did not differ significantly between the two forms. T70 was positively correlated with LT, LMA, and SWC, and negatively correlated with stomatal index (SI). Amax showed positive correlations with SD and SI, but not with Dvein. Vein density was marginally correlated with SD, and significantly correlated with SI. Overall, epiphytic orchids exhibited substantial ecophysiological differentiations from terrestrial species, with the former type showing trait values indicative of greater drought tolerance and increased water storage capacity. The ability to retain water in the leaves plays a key role in maintaining a water balance in those epiphytes. Therefore, the process of transpiration depends less upon the current substrate water supply and enables epiphytic Cymbidium species to adapt more easily to canopy habitats.