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A common dynamical paradigm is that turbulence in the upper ocean is dominated by three classes of motion: mesoscale geostrophic eddies, internal waves and microscale three‐dimensional turbulence. Close to the ocean surface, however, a fourth class of turbulent motion is important: submesoscale frontal dynamics. These have a horizontal scale of O(1–10) km, a vertical scale of O(100) m, and a time scale of O(1) day. Here we review the physical‐chemical‐biological dynamics of submesoscale features, and discuss strategies for sampling them. Submesoscale fronts arise dynamically through nonlinear instabilities of the mesoscale currents. They are ephemeral, lasting only a few days after they are formed. Strong submesoscale vertical velocities can drive episodic nutrient pulses to the euphotic zone, and subduct organic carbon into the ocean's interior. The reduction of vertical mixing at submesoscale fronts can locally increase the mean time that photosynthetic organisms spend in the well‐lit euphotic layer and promote primary production. Horizontal stirring can create intense patchiness in planktonic species. Submesoscale dynamics therefore can change not only primary and export production, but also the structure and the functioning of the planktonic ecosystem. Because of their short time and space scales, sampling of submesoscale features requires new technologies and approaches. This paper presents a critical overview of current knowledge to focus attention and hopefully interest on the pressing scientific questions concerning these dynamics. Key Points Submesoscale physics control ecology locally, but also feedback to basin scales Strong gradients in community structure are created at the submesoscale Despite recent innovations, sampling the submesoscale remains a major challenge

Maximum and minimum stomatal conductance, as well as stomatal size and rate of response, are known to vary widely across plant species, but the functional relationship between these static and dynamic stomatal properties is unknown. The objective of this study was to test three hypotheses: (i) operating stomatal conductance under standard conditions (g op) correlates with minimum stomatal conductance prior to morning light [g min(dawn)]; (ii) stomatal size (S) is negatively correlated with g op and the maximum rate of stomatal opening in response to light, (dg/dt)max; and (iii) g op correlates negatively with instantaneous water-use efficiency (WUE) despite positive correlations with maximum rate of carboxylation (Vc max) and light-saturated rate of electron transport (J max). Using five closely related species of the genus Banksia, the above variables were measured, and it was found that all three hypotheses were supported by the results. Overall, this indicates that leaves built for higher rates of gas exchange have smaller stomata and faster dynamic characteristics. With the aid of a stomatal control model, it is demonstrated that higher g op can potentially expose plants to larger tissue water potential gradients, and that faster stomatal response times can help offset this risk.

From microbes to large predators, there is increasing evidence that marine life is shaped by short-lived submesoscales currents that are difficult to observe, model, and explain theoretically. Whether and how these intense three-dimensional currents structure the productivity and diversity of marine ecosystems is a subject of active debate. Our synthesis of observations and models suggests that the shallow penetration of submesoscale vertical currents might limit their impact on productivity, though ecological interactions at the submesoscale may be important in structuring oceanic biodiversity. Short-lived three-dimensional submesoscale currents, responsible for swirling ocean color chlorophyll filaments, have long been thought to affect productivity. Current research suggests they may not be effective in enhancing phytoplankton growth, but may have important contributions to biodiversity.

Stomatal pores are microscopic structures on the epidermis of leaves formed by 2 specialized guard cells that control the exchange of water vapor and CO₂ between plants and the atmosphere. Stomatal size (S) and density (D) determine maximum leaf diffusive (stomatal) conductance of CO₂ (gcmax) to sites of assimilation. Although large variations in D observed in the fossil record have been correlated with atmospheric CO₂, the crucial significance of similarly large variations in S has been overlooked. Here, we use physical diffusion theory to explain why large changes in S necessarily accompanied the changes in D and atmospheric CO₂ over the last 400 million years. In particular, we show that high densities of small stomata are the only way to attain the highest gcmax values required to counter CO₂\"starvation\" at low atmospheric CO₂ concentrations. This explains cycles of increasing D and decreasing S evident in the fossil history of stomata under the CO₂ impoverished atmospheres of the Permo-Carboniferous and Cenozoic glaciations. The pattern was reversed under rising atmospheric CO₂ regimes. Selection for small S was crucial for attaining high gcmax under falling atmospheric CO₂ and, therefore, may represent a mechanism linking CO₂ and the increasing gas-exchange capacity of land plants over geologic time.

Improvement in crop water-use efficiency (WUE) is a critical priority for regions facing increased drought or diminished groundwater resources. Despite new tools for the manipulation of stomatal development, the engineering of plants with high WUE remains a challenge. We used Arabidopsis epidermal patterning factor (EPF) mutants exhibiting altered stomatal density to test whether WUE could be improved directly by manipulation of the genes controlling stomatal density. Specifically, we tested whether constitutive overexpression of EPF2 reduced stomatal density and maximum stomatal conductance (g w(max)) sufficiently to increase WUE. We found that a reduction in g w(max) via reduced stomatal density in EPF2-overexpressing plants (EPF2OE) increased both instantaneous and long-term WUE without altering significantly the photosynthetic capacity. Conversely, plants lacking both EPF1 and EPF2 expression (epf1epf2) exhibited higher stomatal density, higher g w(max) and lower instantaneous WUE, as well as lower (but not significantly so) long-term WUE. Targeted genetic modification of stomatal conductance, such as in EPF2OE, is a viable approach for the engineering of higher WUE in crops, particularly in future high-carbon-dioxide (CO2) atmospheres.

Blue whales need to time their migration from their breeding grounds to their feeding grounds to avoid missing peak prey abundances, but the cues they use for this are unknown. We examine migration timing (inferred from the local onset and cessation of blue whale calls recorded on seafloor-mounted hydrophones), environmental conditions (e.g., sea surface temperature anomalies and chlorophyll a ), and prey (spring krill biomass from annual net tow surveys) during a 10 year period (2008–2017) in waters of the Southern California Region where blue whales feed in the summer. Colder sea surface temperature anomalies the previous season were correlated with greater krill biomass the following year, and earlier arrival by blue whales. Our results demonstrate a plastic response of blue whales to interannual variability and the importance of krill as a driving force behind migration timing. A decadal-scale increase in temperature due to climate change has led to blue whales extending their overall time in Southern California. By the end of our 10-year study, whales were arriving at the feeding grounds more than one month earlier, while their departure date did not change. Conservation strategies will need to account for increased anthropogenic threats resulting from longer times at the feeding grounds.

To investigate the impact of manipulating stomatal density, a collection of Arabidopsis epidermal patterning factor (EPF) mutants with an approximately 16-fold range of stomatal densities (approx. 20—325% of that of control plants) were grown at three atmospheric carbon dioxide (CO 2 ) concentrations (200, 450 and 1000 ppm), and 30 per cent or 70 per cent soil water content. A strong negative correlation between stomatal size (S) and stomatal density (D) was observed, suggesting that factors that control D also affect S. Under some but not all conditions, mutant plants exhibited abnormal stomatal density responses to CO 2 concentration, suggesting that the EPF signalling pathway may play a role in the environmental adjustment of D. In response to reduced water availability, maximal stomatal conductance was adjusted through reductions in S, rather than D. Plant size negatively correlated with D. For example, at 450 ppm CO 2 EPF2-overexpressing plants, with reduced D, had larger leaves and increased dry weight in comparison with controls. The growth of these plants was also less adversely affected by reduced water availability than plants with higher D, indicating that plants with low D may be well suited to growth under predicted future atmospheric CO 2 environments and/or water-scarce environments.

Given that stomatal movement is ultimately a mechanical process and that stomata are morphologically and mechanically diverse, we explored the influence of stomatal mechanical diversity on leaf gas exchange and considered some of the constraints. Mechanical measurements were conducted on the guard cells of four different species exhibiting different stomatal morphologies, including three variants on the classical \"kidney\" form and one \"dumb-bell\" type; this information, together with gas-exchange measurements, was used to model and compare their respective operational characteristics. Based on evidence from scanning electron microscope images of cryo-sectioned leaves that were sampled under full sun and high humidity and from pressure probe measurements of the stomatal aperture versus guard cell turgor relationship at maximum and zero epidermal turgor, it was concluded that maximum stomatal apertures (and maximum leaf diffusive conductance) could not be obtained in at least one of the species (the grass Triticum aestivum) without a substantial reduction in subsidiary cell osmotic (and hence turgor) pressure during stomatal opening to overcome the large mechanical advantage of subsidiary cells. A mechanism for this is proposed, with a corollary being greatly accelerated stomatal opening and closure. Gas-exchange measurements on T. aestivum revealed the capability of very rapid stomatal movements, which may be explained by the unique morphology and mechanics of its dumb-bell-shaped stomata coupled with \"see-sawing\" of osmotic and turgor pressure between guard and subsidiary cells during stomatal opening or closure. Such properties might underlie the success of grasses.

A long-standing research focus in phytology has been to understand how plants allocate leaf epidermal space to stomata in order to achieve an economic balance between the plant's carbon needs and water use. Here, we present a quantitative theoretical framework to predict allometric relationships between morphological stomatal traits in relation to leaf gas exchange and the required allocation of epidermal area to stomata. Our theoretical framework was derived from first principles of diffusion and geometry based on the hypothesis that selection for higher anatomical maximum stomatal conductance (g smax) involves a trade-off to minimize the fraction of the epidermis that is allocated to stomata. Predicted allometric relationships between stomatal traits were tested with a comprehensive compilation of published and unpublished data on 1057 species from all major clades. In support of our theoretical framework, stomatal traits of this phylogenetically diverse sample reflect spatially optimal allometry that minimizes investment in the allocation of epidermal area when plants evolve towards higher g smax. Our results specifically highlight that the stomatal morphology of angiosperms evolved along spatially optimal allometric relationships. We propose that the resulting wide range of viable stomatal trait combinations equips angiosperms with developmental and evolutionary flexibility in leaf gas exchange unrivalled by gymnosperms and pteridophytes.