Summary: Photosynthesis is currently a focus for crop improvement. The majority of this work has taken place and been assessed in leaves, and limited consideration has been given to the contribution that other green tissues make to whole‐plant carbon assimilation. The major focus of this review is to evaluate the impact of non‐foliar photosynthesis on carbon‐use efficiency and total assimilation. Here we appraise and summarize past and current literature on the substantial contribution of different photosynthetically active organs and tissues to productivity in a variety of different plant types, with an emphasis on fruit and cereal crops. Previous studies provide evidence that non‐leaf photosynthesis could be an unexploited potential target for crop improvement. We also briefly examine the role of stomata in non‐foliar tissues, gas exchange, maintenance of optimal temperatures and thus photosynthesis. In the final section, we discuss possible opportunities to manipulate these processes and provide evidence that Triticum aestivum (wheat) plants genetically manipulated to increase leaf photosynthesis also displayed higher rates of ear assimilation, which translated to increased grain yield. By understanding these processes, we can start to provide insights into manipulating non‐foliar photosynthesis and stomatal behaviour to identify novel targets for exploitation in continuing breeding programmes.

Epidermal Patterning Factor Like 9 (EPFL9), also known as STOMAGEN, is a cysteine-rich peptide that induces stomata formation in vascular plants, acting antagonistically to other epidermal patterning factors (EPF1, EPF2). In grapevine there are two EPFL9 genes, EPFL9-1 and EPFL9-2 sharing 82% identity at protein level in the mature functional C-terminal domain. In this study, CRISPR/Cas9 system was applied to functionally characterize VvEPFL9-1 in ‘Sugraone’, a highly transformable genotype. A set of plants, regenerated after gene transfer in embryogenic calli via Agrobacterium tumefaciens, were selected for evaluation. For many lines, the editing profile in the target site displayed a range of mutations mainly causing frameshift in the coding sequence or affecting the second cysteine residue. The analysis of stomata density revealed that in edited plants the number of stomata was significantly reduced compared to control, demonstrating for the first time the role of EPFL9 in a perennial fruit crop. Three edited lines were then assessed for growth, photosynthesis, stomatal conductance, and water use efficiency in experiments carried out at different environmental conditions. Intrinsic water-use efficiency was improved in edited lines compared to control, indicating possible advantages in reducing stomatal density under future environmental drier scenarios. Our results show the potential of manipulating stomatal density for optimizing grapevine adaptation under changing climate conditions.

Stomata control CO₂ uptake for photosynthesis and water loss through transpiration, thus playing a key role in leaf thermoregulation, water-use efficiency (_iWUE) and plant productivity. In this work, we investigated the relationship between several leaf traits and hypothesized that stomatal behavior to fast (i.e. minutes) environmental changes co-determines along with steady-state traits the physiological response of grapevine to the surrounding fluctuating environment over the growing season. No relationship between _iWUE, heat stress (HS) tolerance and stomatal traits was observed in field grown grapevine, suggesting that other physiological mechanisms are involved in determining leaf evaporative cooling capacity and the seasonal ratio of CO₂ uptake (A) to stomatal conductance (g_s). Indeed, cultivars that in the field had an unexpected combination of high _iWUE but low sensitivity to thermal stress, displayed a quick stomatal closure to light, but a sluggish closure to increased vapor pressure deficit (VPD) levels. This strategy aiming both at conserving water under a high-to-low light transition and in prioritizing evaporative cooling under a low-to-high VPD transition, was mainly observed in Regina and Syrah. Moreover, cultivars with different known responses to soil moisture deficit or high air VPD (isohydric vs anisohydric) had opposite behavior under fluctuating environments, with the isohydric cultivar showing slow stomatal closure to reduced light intensity but quick temporal responses to VPD manipulation. We propose that stomatal behavior to fast environmental fluctuations can play a critical role on leaf thermoregulation and water conservation under natural field conditions in grapevine.

Oilseed rape (Brassica napus L.) yield is strongly decreased by water deficit, and crop-management solutions are urgently required considering the emerging difficulties in breeding for drought-tolerant varieties. Film-forming antitranspirants (polymers) are agrochemicals that, applied to the crop canopy, mechanically block the stomata and decrease canopy transpiration. In this study, the drought-protection efficacy of an adaxial-surface application at the flowering stage of two film-forming treatments (poly-1-p-menthene and di-1-p-menthene) was investigated in pot-grown, droughted oilseed rape over two glasshouse experiments. Over the drought period, the two compounds reduced leaf stomatal conductance (P < 0.001), and as the soil moisture deficit increased, they sustained carbon assimilation and improved water-use efficiency with differing efficacy. Following the antitranspirant treatments, ABA concentration in leaves and reproductive organs was severely reduced and this was accompanied by significant improvements in leaf and flower–pod water potential. Drought significantly decreased the seed dry matter production of oilseed rape plants, by 39% on average. The treatments significantly increased seed dry matter by 13% (poly-1-p-menthene) and 17% (di-1-p-menthene), on average, compared with the unsprayed droughted plants, as a result of a significant increase in number of pods per plant, by 11% and 13%, respectively. The results suggest that film-forming compounds may be a useful crop-management tool to avoid severe drought-induced yield losses in oilseed rape by improving water-use efficiency and plant water status, thus alleviating ABA signalling under water deficit.

The wine sector, among the most profitable agricultural segments, has been markedly affected by the ongoing climate change impacts, such as warmer climate conditions with higher frequency of extreme temperatures and a trend of decreasing precipitation. All this results in higher evaporative demand and therefore higher occurrence of water stress events leading to advancement of temperature-sensitive phenological stages (e.g., budburst and ripening). Such negative effects eventually affect berry development and quality, especially in historically valuable viticultural areas, forcing winegrowers to work within a compressed harvest period to maintain wine typicity. In this work we examined the relationship between environmental variables (air and soil temperature, relative humidity, precipitation, and solar radiation), phenology, berry, and wine quality for the two varieties (Chardonnay and Teroldego) in Trentino Alto-Adige/South Tyrol (Italy) over 36 years. Huglin Index (a bioclimatic heat index), growing degree days (measure of heat accumulation), and overall mean temperature showed linear increase (p < 0.001) in the last years, while no variations were recorded for precipitations. Despite no major effects being observed for phenological interval lengths, the onset of most of the phenological stages for both varieties had significantly (p < 0.001) advanced. However, i) early budburst pushed the budburst-flowering interphase by -1.2 days every two years toward putative colder periods with increased late frost probability and potential slower phenological progression towards flowering, and ii) early veraison shifted the veraison-ripening interphase by 0.25 day per year into warmer periods that oppositely impose faster phenological advancement. Hence, a substantial equilibrium in the seasonal growing length over years was maintained. Potential carry-over effects from the previous season were observed, particularly associated with heat requirements to unlock early phenological events, raising additional concerns on the additive effects of climate change to viticulture. Generally, white wine quality increased (p < 0.05) over the years, while red and sparkling wines remained unaffected. This was putatively related to accurate harvest date decision-making dictated by berry quality parameters: sugar-to-acidity ratio for Chardonnay and bunch sanitary status for Teroldego. Overall, this work provides evidence of the dynamics involved in climate change, and, to our knowledge, its overlooked effects on viticulture, thus providing new insights that can contribute to further developing adaptive strategies.