Both leaf attributes and stomatal traits are linked to water economy in land plants. However, it is unclear whether these two components are associated evolutionarily. In characterizing the possible effect of phylogeny on leaf attributes and stomatal traits, we hypothesized that a correlated evolution exists between the two. Using a phylogenetic comparative method, we analyzed 14 leaf attributes and stomatal traits for 17 species in Paphiopedilum. Stomatal length (SL), stomatal area (SA), upper cuticular thickness (UCT), and total cuticular thickness (TCT) showed strong phylogenetic conservatism whereas stomatal density (SD) and stomatal index (SI) were significantly convergent. Leaf vein density was correlated with SL and SD whether or not phylogeny was considered. The lower epidermal thickness (LET) was correlated positively with SL, SA, and stomatal width but negatively with SD when phylogeny was not considered. When this phylogenetic influence was factored in, only the significant correlation between SL and LET remained. Our results support the hypothesis for correlated evolution between stomatal traits and vein density in Paphiopedilum. However, they do not provide evidence for an evolutionary association between stomata and leaf thickness. These findings lend insight into the evolution of traits related to water economy for orchids under natural selection.

Drought stress can induce closure of stomata, thus leading to photoinhibition. The effects of prolonged severe drought under natural growing conditions on photosystem I (PSI), photosystem II (PSII) and cyclic electron flow (CEF) in drought-tolerant tree species are unclear. In spring 2010, southwestern China confronted severe drought that lasted several months. Using three dominant evergreen species, Cleistanthus sumatranus (Miq.) Muell. Arg. (Euphorbiaceae), Celtis philippensis Bl. (Ulmaceae) and Pistacia weinmannifolia J. Poisson ex Franch. (Anacardiaceae) that are native to a tropical limestone forest, we investigated the influence of this stress on PSI and PSII activities as well as light energy distribution in the PSII and P700 redox state. By the end of the drought period, predawn leaf water potential (Ψ_pd) largely declined in each species, especially in C. sumatranus. Photosystem I activity strongly decreased in the three species, especially in C. sumatranus which showed a decrease of 65%. The maximum quantum yield of PSII after dark adaptation remained stable in P. weinmannifolia and C. philippensis but significantly decreased in C. sumatranus. Light response curves indicated that both linear electron flow and non-photochemical quenching were severely inhibited in C. sumatranus along with disappearance of CEF, resulting in deleterious excess light energy in PSII. We conclude that PSI is more sensitive than PSII to prolonged severe drought in these three drought-tolerant species, and CEF is essential for photoprotection in them.

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.

Floral longevity (FL) determines the balance between pollination success and flower maintenance. While a longer floral duration enhances the ability of plants to attract pollinators, it can be detrimental if it negatively affects overall plant fitness. Longer-lived leaves display a positive correlation with their dry mass per unit area, which influences leaf construction costs and physiological functions. However, little is known about the association among FL and floral dry mass per unit area (FMA) and water maintenance traits. We investigated whether increased FL might incur similar costs. Our assessment of 11 species of Paphiopedilum (slipper orchids) considered the impact of FMA and flower water-maintenance characteristics on FL. We found a positive relationship between FL and FMA. Floral longevity showed significant correlations with osmotic potential at the turgor loss and bulk modulus of elasticity but not with FA. Neither the size nor the mass per area was correlated between leaves and flowers, indicating that flower and leaf economic traits evolved independently. Therefore, our findings demonstrate a clear relationship between FL and the capacity to maintain water status in the flower. These economic constraints also indicate that extending the flower life span can have a high physiological cost in Paphiopedilum.