Ear photosynthesis in C3 cereals and its contribution to grain yield: methodologies, controversies, and perspectives.

In C3 cereals such as wheat and barley, grain filling was traditionally explained as being sustained by assimilates from concurrent leaf photosynthesis and remobilization from the stem. In recent decades, a role for ear photosynthesis as a contributor to grain filling has emerged. This review analyzes several aspects of this topic: (i) methodological approaches for estimation of ear photosynthetic contribution to grain filling; (ii) the existence of genetic variability in the contribution of the ear, and evidence of genetic gains in the past; (iii) the controversy of the existence of C4 metabolism in the ear; (iv) the response of ear photosynthesis to water deficit; and (v) morphological and physiological traits possibly related to ear temperature and thermal balance of the ear. The main conclusions are: (i) there are a number of methodologies to quantify ear photosynthetic activity (e.g. gas exchange and chlorophyll fluorescence) and the contribution of the ear to grain filling (individual ear shading, ear emergence in shaded canopies, and isotope composition); (ii) the contribution of ear photosynthesis seems to have increased in modern wheat germplasm; (iii) the contribution of the ear to grain filling increases under resource-limitation (water deficit, defoliation, or pathogen infection); (iv) there is genetic variability in the contribution of the ear in wheat, opening up the possibility to use this trait to ameliorate grain yield; (v) current evidence supports the existence of C3 metabolism rather than C4 metabolism; (vi) the ear is a 'dehydration avoider organ' under drought; and (vii) thermal balance in the ear is a relevant issue to explore, and more research is needed to clarify the underlying morphological and physiological traits.

Frequently asked questions about chlorophyll fluorescence, the sequel.

Using chlorophyll (Chl) a fluorescence many aspects of the photosynthetic apparatus can be studied, both in vitro and, noninvasively, in vivo. Complementary techniques can help to interpret changes in the Chl a fluorescence kinetics. Kalaji et al. (Photosynth Res 122:121-158, 2014a) addressed several questions about instruments, methods and applications based on Chl a fluorescence. Here, additional Chl a fluorescence-related topics are discussed again in a question and answer format. Examples are the effect of connectivity on photochemical quenching, the correction of F V /F M values for PSI fluorescence, the energy partitioning concept, the interpretation of the complementary area, probing the donor side of PSII, the assignment of bands of 77 K fluorescence emission spectra to fluorescence emitters, the relationship between prompt and delayed fluorescence, potential problems when sampling tree canopies, the use of fluorescence parameters in QTL studies, the use of Chl a fluorescence in biosensor applications and the application of neural network approaches for the analysis of fluorescence measurements. The answers draw on knowledge from different Chl a fluorescence analysis domains, yielding in several cases new insights.

Frequently asked questions about in vivo chlorophyll fluorescence: practical issues.

The aim of this educational review is to provide practical information on the hardware, methodology, and the hands on application of chlorophyll (Chl) a fluorescence technology. We present the paper in a question and answer format like frequently asked questions. Although nearly all information on the application of Chl a fluorescence can be found in the literature, it is not always easily accessible. This paper is primarily aimed at scientists who have some experience with the application of Chl a fluorescence but are still in the process of discovering what it all means and how it can be used. Topics discussed are (among other things) the kind of information that can be obtained using different fluorescence techniques, the interpretation of Chl a fluorescence signals, specific applications of these techniques, and practical advice on different subjects, such as on the length of dark adaptation before measurement of the Chl a fluorescence transient. The paper also provides the physiological background for some of the applied procedures. It also serves as a source of reference for experienced scientists.

Glyphosate effects on gas exchange and chlorophyll fluorescence responses of two Lolium perenne L. biotypes with differential herbicide sensitivity.

Despite the extensive use of glyphosate, how it alters the physiology and metabolism of plants is still unclear. Photosynthesis is not regarded to be a primary inhibitory target of glyphosate, but it has been reported to be affected by this herbicide. The aim of the current research was to determine the effects of glyphosate on the light and dark reactions of photosynthesis by comparing glyphosate-susceptible and glyphosate-resistant Lolium perenne biotypes. After glyphosate treatment, accumulation of reduced carbohydrates occurred before a decrease in gas exchange. Stomatal conductance and CO(2) assimilation were reduced earlier than chlorophyll fluorescence and the amount of chlorophyll in susceptible plants. In the glyphosate-resistant biotype, stomatal conductance was the only parameter slightly affected only 5 days post-application. In susceptible plants, the initial glyphosate effects on gas exchange could be a response to a feedback regulation of photosynthesis. Since the herbicide affects actively growing tissues regardless of the inhibition of photosynthesis, the demand of assimilates decreased and consequently induced an accumulation of carbohydrates in leaves. We concluded that stomatal conductance could be a very sensitive parameter to assess both the susceptibility/resistance to glyphosate before the phytotoxic symptoms become evident.

Control of ascorbic acid synthesis and accumulation and glutathione by the incident light red/far red ratio in Phaseolus vulgaris leaves.

The effects of red/far red (R/FR) ratios on leaf ascorbate (AA) and glutathione (GSH) accumulation were examined in common bean (Phaseolus vulgaris L.). Growth under low R/FR ratios resulted in a "shade" phenotype and much lower leaf AA and GSH contents than high (R/FR) ratios. Photosynthesis rates were unaffected by changes in the R/FR ratio but leaf respiration rates, pyridine nucleotide pools and antioxidant enzyme activities were decreased under the low R/FR regime. The GSH pool changed slowly in response to altered R/FR ratios but leaf ascorbate acclimated over a single photoperiod. We conclude that light quality signals, particularly R/FR ratios, are important regulators of antioxidant synthesis and accumulation. These acclimatory changes are an early response to changing light environment.

Ear of durum wheat under water stress: water relations and photosynthetic metabolism.

The photosynthetic characteristics of the ear and flag leaf of well-watered (WW) and water-stressed (WS) durum wheat (Triticum turgidum L. var. durum) were studied in plants grown under greenhouse and Mediterranean field conditions. Gas exchange measurements simultaneously with modulated chlorophyll fluorescence were used to study the response of the ear and flag leaf to CO2 and O2 during photosynthesis. C4 metabolism was identified by assessing the sensitivity of photosynthetic rate and electron transport to oxygen. The presence of CAM metabolism was assessed by measuring daily patterns of stomatal conductance and net CO2 assimilation. In addition, the histological distribution of Rubisco protein in the ear parts was studied by immunocytochemical localisation. Relative water content (RWC) and osmotic adjustment (osmotic potential at full turgor) were also measured in these organs. Oxygen sensitivity of the assimilation rate and electron transport, the lack of Rubisco compartmentalisation in the mesophyll tissues and the gas-exchange pattern at night indicated that neither C4 nor CAM metabolism occurs in the ear of WW or WS plants. Nevertheless, photosynthetic activity of the flag leaf was more affected by WS conditions than that of the ear, under both growing conditions. The lower sensitivity under water stress of the ear than of the flag leaf was linked to higher RWC and osmotic adjustment in the ear bracts and awns. We demonstrate that the better performance of the ear under water stress (compared to the flag leaf) is not related to C4 or CAM photosynthesis. Rather, drought tolerance of the ear is explained by its higher RWC in drought. Osmotic adjustment and xeromorphic traits of ear parts may be responsible.

Photoprotection in water-stressed plants of durum wheat (Triticum turgidum var. durum): changes in chlorophyll fluorescence, spectral signature and photosynthetic pigments.

We analysed the photoprotective response in water-stressed plants of durum wheat (Triticum turgidum L. var. durum cv. Mexa). The plants were grown in a greenhouse for 4 weeks and then exposed to water stress by withholding water for 8 d. Development of water stress was monitored as the decrease in relative water content (RWC) and net CO2 assimilation of the last fully developed leaf. The photoprotective response was evaluated in the same leaves by analysing modulated chlorophyll fluorescence, leaf spectroradiometrical changes, and pigment content. Measurements were performed 3, 6 (moderate stress) and 8 (severe stress) d after water-stress treatment began. The non-photochemical quenching of chlorophyll fluorescence (qN), as well as the contents of zeaxanthin and antheraxanthin increased significantly after 6 d of treatment. However, a further rise in these xanthophylls on day 8 was not associated with any increase in qN. In addition, the ß-carotene content rose significantly on day 8, suggesting an increase in antioxidant defences. The photochemical index (PI), derived from spectroradiometrical measurements, showed a strong progressive drop on days 6 and 8, which was paralleled by an increase in the de-epoxidation state of the xanthophyll cycle (DPS), in particular by the zeaxanthin content. At midday, PI was strongly (negatively) correlated with DPS and qN. These results suggest that the PI may be a reliable indicator of photoprotection in the study of plant stress, and in breeding programs.