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Crop domestication for increasing growth rates and yields appears to have altered the features of adaxial and abaxial stomata, but its effect on leaf water use efficiency (WUE) have not been experimentally verified. In this study, we characterized stomatal anatomy and carbon isotope discrimination (δ13C) in 32 wild and 36 domesticated genotypes of cotton grown under agricultural field conditions. The results showed that domesticated genotypes possessed lower WUE, as indicated by low or more negative δ13C compared with wild genotypes. Higher theoretical maximum stomatal conductance (gsmax) after domestication was accounted for by more stomata rather than significantly enlarged stomata. Specifically, abaxial stomatal density was higher whilst there was no change in the adaxial density. The size of both adaxial and abaxial stomata was greater due to larger guard cells but without there being any increase in pore size. However, there was a negative relationship between δ13C and stomatal size across wild and domesticated genotypes, especially on the abaxial leaf surface, because bigger stomata resulted in a lower maximum stomatal response rate to fluctuating canopy light, resulting in increased water loss. Overall, our results indicate that cotton domestication has resulted in substantial variation in stomatal anatomy, and that WUE and drought tolerance can potentially be improved in future breeding by decreasing the size of abaxial stomata to produce a faster stomatal response and hence a reduction in unnecessary water loss.

Background and Aims The process of domestication has driven dramatic shifts in plant functional traits, including leaf mass per area (LMA). It remains unclear whether domestication has produced concerted shifts in the lower-level anatomical traits that underpin LMA and how these traits in turn affect photosynthesis. Methods In this study we investigated controls of LMA and leaf gas exchange by leaf anatomical properties at the cellular, tissue and whole-leaf levels, comparing 26 wild and 31 domesticated genotypes of cotton (Gossypium). Key Results As expected, domesticated plants expressed lower LMA, higher photosynthesis and higher stomatal conductance, suggesting a shift towards the ‘faster’ end of the leaf economics spectrum. At whole-leaf level, variation in LMA was predominantly determined by leaf density (LD) both in wild and domesticated genotypes. At tissue level, higher leaf volume per area (Vleaf) in domesticated genotypes was driven by a simultaneous increase in the volume of epidermal, mesophyll and vascular bundle tissue and airspace, while lower LD resulted from a lower volume of palisade tissue and vascular bundles (which are of high density), paired with a greater volume of epidermis and airspace, which are of low density. The volume of spongy mesophyll exerted direct control on photosynthesis in domesticated genotypes but only indirect control in wild genotypes. At cellular level, a shift to larger but less numerous cells with thinner cell walls underpinned a lower proportion of cell wall mass, and thus a reduction in LD. Conclusions Taken together, cotton domestication has triggered synergistic shifts in the underlying determinants of LMA but also photosynthesis, at cell, tissue and whole-leaf levels, resulting in a marked shift in plant ecological strategy.

Background and Aims The leaf economic spectrum (LES) is related to dry mass and nutrient investments towards photosynthetic processes and leaf structures, and to the duration of returns on those investments (leaf lifespan, LL). Phosphorus (P) is a key limiting nutrient for plant growth, yet it is unclear how the allocation of leaf P among different functions is coordinated with the LES. We addressed this question among 12 evergreen woody species co-occurring on P-impoverished soils in south-eastern Australia. Methods Leaf ‘economic’ traits, including LL, leaf mass per area (LMA), light-saturated net photosynthetic rate per mass (Amass), dark respiration rate, P concentration ([Ptotal]), nitrogen concentration, and P resorption, were measured for three pioneer and nine non-pioneer species. Leaf P was separated into five functional fractions: orthophosphate P (Pi), metabolite P (PM), nucleic acid P (PN), lipid P (PL), and residual P (PR; phosphorylated proteins and unidentified compounds that contain P). Results LL was negatively correlated with Amass and positively correlated with LMA, representing the LES. Pioneers occurred towards the short-LL end of the spectrum and exhibited higher [Ptotal] than non-pioneer species, primarily associated with higher concentrations of Pi, PN and PL. There were no significant correlations between leaf P fractions and LL or LMA, while Amass was positively correlated with the concentration of PR. Conclusions Allocation of leaf P to different fractions varied substantially among species. This variation was partially associated with the LES, which may provide a mechanism underlying co-occurrence of species with different ecological strategies under P limitation.

Domestication involves dramatic phenotypic and physiological diversifications due to successive selection by breeders toward high yield and quality. Although photosynthetic nitrogen use efficiency (PNUE) is a major trait for understanding leaf nitrogen economy, it is unclear whether PNUE of cotton has been improved under domestication. Here, we investigated the effect of domestication on nitrogen allocation to photosynthetic machinery and PNUE in 25 wild and 37 domesticated cotton genotypes. The results showed that domesticated genotypes had higher nitrogen content per mass (Nm), net photosynthesis under saturated light (Asat), and PNUE but similar nitrogen content per area (Na) compared with wild genotypes. As expected, in both genotypes, PNUE was positively related to Asat but negatively correlated with Na. However, the relative contribution of Asat to PNUE was greater than the contribution from Na. Domesticated genotypes had higher nitrogen allocation to light-harvesting (NL, nitrogen in light-harvesting chlorophyll-protein complex), to bioenergetics (Nb, total nitrogen of cytochrome f, ferredoxin NADP reductase, and the coupling factor), and to Rubisco (Nr) than wild genotypes; however, the two genotype groups did not differ in PNUEp, the ratio of Asat to Np (itself the sum of NL, Nb, and Nr). Our results suggest that more nitrogen allocation to photosynthetic machinery has boosted Asat under cotton domestication. Improving the efficiency of nitrogen use in photosynthetic machinery might be future aim to enhance Asat of cotton.

Mesophyll conductance (gm) is a crucial leaf trait contributing to the photosynthetic rate (AN). Plant domestication typically leads to an enhancement of AN that is often associated with profound anatomical modifications, but it is unclear which of these structural alterations influence gm. We analyzed the implication of domestication on leaf anatomy and its effect on gm in 26 wild and 31 domesticated cotton genotypes (Gossypium sp.) grown under field conditions. We found that domesticated genotypes had higher AN but similar gm to wild genotypes. Consistent with this, domestication did not translate into significant differences in the fraction of mesophyll occupied by intercellular air spaces (fias) or mesophyll and chloroplast surface area exposed to intercellular air space (Sm/S and Sc/S, respectively). However, leaves of domesticated genotypes were significantly thicker, with larger but fewer mesophyll cells with thinner cell walls. Moreover, domesticated genotypes had higher cell wall conductance (gcw) but smaller cytoplasmic conductance (gcyt) than wild genotypes. It appears that domestication in cotton has not generally led to significant improvement in gm, in part because their thinner mesophyll cell walls (increasing gcw) compensate for their lower gcyt, itself due to larger distance between plasmalemma and chloroplast envelopes.

Crop domestication appears to alter the adaxial and abaxial stomatal features for increasing growth rate and yield. However, its effect on leaf water use efficiency (WUE) has not been experimentally verified under domestication. In this study, we characterized stomatal anatomy and carbon isotope (δ 13C) in 32 wild and 36 domesticated genotypes of cotton grown under agricultural field. The results showed that domesticated genotypes possessed lower WUE as indicated by low or more negative δ 13C compared with wild genotypes. Higher maximum theoretical stomatal conductance (gsmax) was underpinned by more stomata rather than significantly enlarged stomata after domestication. Specifically, abaxial stomatal density was higher but no change in adaxial stomatal density following domestication, while both adaxial and abaxial stomatal size were greater due to larger guard cell without the contribution to stomatal pore size. However, there was a negative relationship between δ 13C and SS, especially the abaxial SS across wild and domesticated genotypes, because enlarged stomata resulted in a lower stomatal response rate to fluctuating canopy light, consequently excessive water loss. In conclusion, cotton domestication has triggered substantial variation in stomatal anatomy. WUE and drought tolerance in cotton can be improved by decreasing abaxial stomatal size for faster stomatal response and reducing unnecessary water loss in future cotton breeding.

There is a trade-off between leaf structures and photosynthesis physiology. While numerous reports have shown a dif- ference in photosynthesis between Gossypium hirsutum L. (upland cotton) and G. barbadense L. (pima cotton), the potential contribution of leaf structures on this difference was not fully clarified. Here, we investigated the differences in area- and mass-based photosynthetic traits and leaf structures between upland and pima cottons. Our results showed that upland cotton had higher area-based net photosynthetic rate (Parea) than that of pima cotton, which was attributed to the coordination of stomatal conductance (gs), area-based mesophyll conductance (gm-area), and maximum carboxylation rate (Vcmax-area). Parea, gm-area, and Vcmax-area correlated positively with leaf mass per area. But upland and pima cotton had similar mass-based PN (Pmass), gm (gm-mass), Vcmax (Vcmax-mass), suggesting that they have a similar photosynthetic capacity of single cells. Consequently, the higher area-based values in upland cotton were only due to a higher leaf thickness (Tleaf).

Extreme environments, such as deserts and high-elevation ecosystems, are very important from biodiversity and ecological perspectives. However, plant physiology at those sites has been scarcely studied, likely due to logistic difficulties. In the present study, leaf physiological traits in native plants were analyzed from arid zones across an elevational transect in Western China, from Turpan Basin to the Qinghai-Tibet Plateau (QTP) at Delingha. The aim of this study was to use leaf physiological traits to help identifying potentially threatened species and true extremophiles. Physiological measurements in the field, and particularly in situ measurements of gas exchange and chlorophyll fluorescence, have been determined to be useful to determine the current state of plants at a given environment. Using this approach plus a combination of leaf traits, several species performing particularly well at the QTP were identified, e.g. Hedysarum multijugum, as well as at Manas drylands, e.g. Peganum harmala and Setaria viridis. On the other hand, several species showed marked signs of severe stress, in particular a very low photosynthetic rate over its potential maximum, as well as other negative traits, like low water and/or nitrogen-use-efficiency, which should be considered in conservation plans. Interestingly, all C

Drought stress causes changes in vein and stomatal density. The objectives of this study were to determine (1) if the changes in vein and stomatal density are coordinated in cotton (Gossypium hirsutum L.) and (2) how these changes affect water-use efficiency (WUE). The results showed significant positive correlations between vein density and stomatal density when cotton was grown under different degrees of drought stress. WUE was significantly positively correlated with the densities of both veins and stomata. Stomatal pore area and stomatal density on the abaxial leaf side, but not the adaxial side, were significantly correlated with WUE, stomatal conductance, leaf net photosynthetic rate, and transpiration rate. In conclusion, coordinated changes in vein and stomatal density improve the WUE of cotton under drought stress. The abaxial leaf side plays a more important role than the adaxial side in WUE and gas exchange.

Plant density is a common practice for optimizing canopy photosynthetic capacity and improving cotton ( Gossypium hirsutum L.) yield. The objective of this study was to evaluate the effects of plant density (14.5, 19.5 and 29 plants m −2 were adjusted by wide, medium and narrow row spacing, respectively) on canopy apparent photosynthetic use efficiency of light (CAP LUE ) and N (CAP NUE ), dry matter accumulation and yield formation of cotton plants (cv. SCRC24). Cotton in narrow row spacing responded to light interception and distribution within its canopy by significantly increasing the proportion of leaf area index (LAI) in the upper canopy and by increasing leaf N content per unit land area ( N land ). Cotton in wide row spacing compared with narrow row spacing had a significantly greater proportion of LAI in the mid and lower canopy and a significantly lower N land . The changes in canopy structure and leaf photosynthetic characteristics in response to row spacing affected the relationship between CAP and light within the canopies. Under low light conditions, CAP LUE and CAP NUE were greater in narrow row spacing than in wide row spacing. Under high light conditions, maximum CAP LUE and CAP NUE were greater in wide row spacing than in narrow row spacing. These light response curves support the hypothesis that CAP LUE and CAP NUE were maximized according to the light interception and distribution within its canopy, which is by adjusting canopy structure and leaf photosynthetic characteristics. Furthermore, there was no difference in leaf + boll dry weight among the three row spacing treatments. The leaf + boll dry weight was maintained in narrow row spacing by increasing leaf dry weight at the expense of boll dry weight. The cotton responded to wide row spacing both by increasing the single boll weight and the proportion of LAI in the mid- and lower canopy and by increasing the proportion of bolls in the upper canopy. Cotton in wide row spacing had better canopy leaf and boll distribution, therefore higher CAP, because a greater proportion of leaf area was allocated to the mid canopy and the canopy structure was optimal. This indicated that yield was influenced not only by maximum CAP, CAP LUE , and CAP NUE , but also by the spatial distribution of vegetative and reproductive organs.

Mesophyll conductance (gm ) is one of the major determinants of photosynthetic rate, for which it has an impact on crop yield. However, the regulatory mechanisms behind the decline in gm of cotton (Gossypium. spp) by drought are unclear. An upland cotton (Gossypium hirsutum) genotype and a pima cotton (Gossypium barbadense) genotype were used to determine the gas exchange parameters, leaf anatomical structure as well as aquaporin and carbonic anhydrase gene expression under well-watered and drought treatment conditions. In this study, the decrease of net photosynthetic rate (AN ) under drought conditions was related to a decline in gm and in stomatal conductance (gs ). gm and gs coordinate with each other to ensure optimum state of CO2 diffusion and achieve the balance of water and CO2 demand in the process of photosynthesis. Meanwhile, mesophyll limitations to photosynthesis are equally important to the stomatal limitations. Considering gm , its decline in cotton leaves under drought was mostly regulated by the chloroplast surface area exposed to leaf intercellular air spaces per leaf area (Sc /S) and might also be regulated by the expression of leaf CARBONIC ANHYDRASE (CA1). Meanwhile, cotton leaves can minimize the decrease in gm under drought by maintaining cell wall thickness (Tcw ). Our results indicated that modification of chloroplasts might be a target trait in future attempts to improve cotton drought tolerance.

Mesophyll conductance (gm) has been shown to affect photosynthetic capacity and thus the estimates of terrestrial carbon balance. While there have been some attempts to model gm at the leaf and larger scales, the potential contribution of gm to the photosynthesis of non-leaf green organs has not been studied. Here, we investigated the influence of gm on photosynthesis of cotton bracts and how it in turn is influenced by anatomical structures, by comparing leaf palisade and spongy mesophyll with bract tissue. Our results showed that photosynthetic capacity in bracts is much lower than in leaves, and that gm is a limiting factor for bract photosynthesis to a similar extent to stomatal conductance. Bract and the spongy tissue of leaves have lower mesophyll conductance than leaf palisade tissue due to the greater volume fraction of intercellular air spaces, smaller chloroplasts, lower surface area of mesophyll cells and chloroplasts exposed to leaf intercellular air spaces and, perhaps, lower membrane permeability. Comparing bracts with leaf spongy tissue, although bracts have a larger cell wall thickness, they have a similar gm estimated from anatomical characteristics, likely due to the cumulative compensatory effects of subtle differences in each subcellular component, especially chloroplast traits. These results provide the first evidence for anatomical constraints on gm and photosynthesis in non-leaf green organs.