In many plant species that remain leafless part of the year, CO 2 fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO 2 fixation by stems with stomata in the epidermis and net corticular CO 2 fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO 2 recycling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO 2 uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO 2 occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO 2 , the latter figure implying net CO 2 uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.
In obligate Crassulacean acid metabolism (CAM), up to 99 % of CO(2) assimilation occurs during the night, therefore supporting the hypothesis that CAM is adaptive because it allows CO(2) fixation during the part of the day with lower evaporative demand, making life in water-limited environments possible. By comparison, in facultative CAM (inducible CAM, C(3)-CAM) and CAM-cycling plants drought-induced dark CO(2) fixation may only be, with few exceptions, a small proportion of C(3) CO(2) assimilation in watered plants and occur during a few days. From the viewpoint of survival the adaptive advantages, i.e. increased fitness, of facultative CAM and CAM-cycling are not obvious. Therefore, it is hypothesized that, if it is to increase fitness, CAM must aid in reproduction. Scope An examination of published reports of 23 facultative CAM and CAM-cycling species finds that, in 19 species, drought-induced dark CO(2) fixation represents on average 11 % of C(3) CO(2) assimilation of watered plants. Evidence is discussed on the impact of the operation of CAM in facultative and CAM-cycling plants on their survival--carbon balance, water conservation, water absorption, photo-protection of the photosynthetic apparatus--and reproductive effort. It is concluded that in some species, but not all, facultative and cycling CAM contribute, rather than to increase carbon balance, to increase water-use efficiency, water absorption, prevention of photoinhibition and reproductive output.
Water relations and photosynthetic characteristics of plants of Lycium nodosum grown under increasing water deficit (WD), saline spray (SS) or saline irrigation (SI) were studied. Plants of this perennial, deciduous shrub growing in the coastal thorn scrubs of Venezuela show succulent leaves which persist for approx. 1 month after the beginning of the dry season; leaf succulence is higher in populations closer to the sea. These observations suggested that L. nodosum is tolerant both to WD and salinity. In the glasshouse, WD caused a marked decrease in the xylem water potential (psi), leaf osmotic potential (psi(s)) and relative water content (RWC) after 21 d; additionally, photosynthetic rate (A), carboxylation efficiency (CE) and stomatal conductance (gs) decreased by more than 90 %. In contrast, in plants treated for 21 d with a foliar spray with 35 per thousand NaCl or irrigation with a 10 % NaCl solution, psi and RWC remained nearly constant, while psi(s) decreased by 30 %, and A, CE and gs decreased by more than 80 %. An osmotic adjustment of 0.60 (SS) and 0.94 MPa (SI) was measured. Relative stomatal and mesophyll limitations to A increased with both WD and SS, but were not determined for SI-treated plants. No evidence of chronic photoinhibition due to any treatment was observed, since maximum quantum yield of PSII, Fv/Fm, did not change with either drought in the field or water or salinity stress in the glasshouse. Nevertheless, WD and SI treatments caused a decrease in the photochemical (qP) and an increase in the non-photochemical (qN) quenching coefficients relative to controls; qN was unaffected by the SS treatment. The occurrence of co-limitation of A by stomatal and non-stomatal factors in plants of L. nodosum may be associated with the extended leaf duration under water or saline stress. Additionally, osmotic adjustment may partly explain the relative maintenance of A and gs in the SS and SI treatments and the tolerance to salinity of plants of this species in coastal habitats.
This review summarizes the research on physiological responses to flooding of trees in the seasonal black-water wetland of the Mapire River in Venezuela. Inter-annual variability was found during 8 years of sampling, in spite of which a general picture emerged of increased stomatal conductance (gs) and photosynthetic rate (PN) during the flooded period to values as high as or higher than in plants in drained wet soil. Models explaining the initial inhibitory responses and the acclimation to flooding are proposed. In the inhibitory phase of flooding, hypoxia generated by flooding causes a decrease in root water absorption and stomatal closure. An increase with flooding in xylem water potential (ψ) suggests that flooding does not cause water deficit. The PN decreases due to changes in relative stomatal and non-stomatal limitations to photosynthesis; an increase in the latter is due to reduced chlorophyll and total soluble protein content. Total non-structural carbohydrates (TNC) accumulate in leaves but their content begins to decrease during the acclimatized phase at full flooding, coinciding with the resumption of high gs and PN. The reversal of the diminution in gs is associated, in some but not all species, to the growth of adventitious roots. The occurrence of morpho-anatomical and biochemical adaptations which improve oxygen supply would cause the acclimation, including increased water absorption by the roots, increased rubisco and chlorophyll contents and ultimately increased PN. Therefore, trees would perform as if flooding did not signify a stress to their physiology.
Cocoa grows under shade, but some cultivars develop successfully in full sunlight. In order to characterize the response to photosynthetic photon flux density (PPFD) of a Modern Criollo cocoa clone, gas exchange, photochemical activity and leaf traits, and their relation to growth were measured in seedlings growing in a greenhouse at three different values of PPFD, as well as in adults in full sunlight and shade in the field. Plants showed changes in physiological, biochemical, and morpho‐anatomical traits in response to the different light conditions, and in the phenotypic plasticity of these variables. Seedlings subjected to high PPFD in the greenhouse showed decreases in photosynthetic rate, apparent quantum yield of CO2 fixation and photochemical quenching, and increases in non‐photochemical quenching, suggesting down‐regulation of PSII. In contrast, trees under full sunlight in the field showed a marked reduction in maximum quantum yield of PSII, indicating photoinhibition and supporting that cocoa is a shade tolerant crop. Cocoa showed higher plasticity of physiological and biochemical variables than morpho‐anatomical variables in response to PPFD. Effects of time under treatment in the greenhouse and plant age (greenhouse vs field) on plasticity were observed. The acclimation observed in some of the variables studied after 6 months in high light did not represent a particular advantage to seedlings, since relative growth rate was lower than in low‐ and medium‐light seedlings.
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