Perennial grains hold promise, especially for marginal landscapes or with limited resources where annual versions struggle.
In open places, plants are exposed to higher fluence rates of photosynthetically active radiation and to higher red to far-red ratios than under the shade of neighbor plants. High fluence rates are known to increase stomata density. Here we show that high, compared to low, red to far-red ratios also increase stomata density in Arabidopsis (Arabidopsis thaliana). High red to farred ratios increase the proportion of phytochrome B (phyB) in its active form and the phyB mutant exhibited a constitutively low stomata density. phyB increased the stomata index (the ratio between stomata and epidermal cells number) and the level of anphistomy (by increasing stomata density more intensively in the adaxial than in the abaxial face). phyB promoted the expression of FAMA and TOO MANY MOUTHS genes involved in the regulation of stomata development in young leaves. Increased stomata density resulted in increased transpiration per unit leaf area. However, phyB promoted photosynthesis rates only at high fluence rates of photosynthetically active radiation. In accordance to these observations, phyB reduced long-term water-use efficiency estimated by the analysis of isotopic discrimination against 13 CO 2 . We propose a model where active phyB promotes stomata differentiation in open places, allowing plants to take advantage of the higher irradiances at the expense of a reduction of water-use efficiency, which is compensated by a reduced leaf area.
Waterlogging is expected to increase as a consequence of global climate change, constraining crop production in various parts of the world. This study assessed tolerance to 14-days of early- or late-stage waterlogging of the major winter crops wheat, barley, rapeseed and field pea. Aerenchyma formation in adventitious roots, leaf physiological parameters (net photosynthesis, stomatal and mesophyll conductances, chlorophyll fluorescence), shoot and root growth during and after waterlogging, and seed production were evaluated. Wheat produced adventitious roots with 20–22% of aerenchyma, photosynthesis was maintained during waterlogging, and seed production was 86 and 71% of controls for early- and late-waterlogging events. In barley and rapeseed, plants were less affected by early- than by late-waterlogging. Barley adventitious roots contained 19% aerenchyma, whereas rapeseed did not form aerenchyma. In barley, photosynthesis was reduced during early-waterlogging mainly by stomatal limitations, and by non-stomatal constraints (lower mesophyll conductance and damage to photosynthetic apparatus as revealed by chlorophyll fluorescence) during late-waterlogging. In rapeseed, photosynthesis was mostly reduced by non-stomatal limitations during early- and late-waterlogging, which also impacted shoot and root growth. Early-waterlogged plants of both barley and rapeseed were able to recover in growth upon drainage, and seed production reached ca. 79–85% of the controls, while late-waterlogged plants only attained 26–32% in seed production. Field pea showed no ability to develop root aerenchyma when waterlogged, and its photosynthesis (and stomatal and mesophyll conductances) was rapidly decreased by the stress. Consequently, waterlogging drastically reduced field pea seed production to 6% of controls both at early- and late-stages with plants being unable to resume growth upon drainage. In conclusion, wheat generates a set of adaptive responses to withstand 14 days of waterlogging, barley and rapeseed can still produce significant yield if transiently waterlogged during early plant stages but are more adversely impacted at the late stage, and field pea is not suitable for areas prone to waterlogging events of 14 days at either growth stage.
While the most conspicuous response to low red/far-red ratios (R:FR) of shade light perceived by phytochrome is the promotion of stem growth, additional, less obvious effects may be discovered by studying changes in the stem transcriptome. Here, we report rapid and reversible stem transcriptome responses to R:FR in tomato (Solanum lycopersicum). As expected, low R:FR promoted the expression of growth-related genes, including those involved in the metabolism of cell wall carbohydrates and in auxin responses. In addition, genes involved in flavonoid synthesis, isoprenoid metabolism, and photosynthesis (dark reactions) were overrepresented in clusters showing reduced expression in the stem of low R:FR-treated plants. Consistent with these responses, low R:FR decreased the levels of flavonoids (anthocyanin, quercetin, kaempferol) and selected isoprenoid derivatives (chlorophyll, carotenoids) in the stem and severely reduced the photosynthetic capacity of this organ. However, lignin contents were unaffected. Low R:FR reduced the stem levels of jasmonate, which is a known inducer of flavonoid synthesis. The rate of stem respiration was also reduced in low R:FR-treated plants, indicating that by downsizing the stem photosynthetic apparatus and the levels of photoprotective pigments under low R:FR, tomato plants reduce the energetic cost of shadeavoidance responses.
Lotus corniculatus L. and Lotus glaber Mill. are warm-season legume species adapted to many kinds of environmental stress, including flooding conditions, whereas other popular forage legumes, like alfalfa or white clover, cannot thrive. This study evaluates the relationship between root aerenchyma, water relations and leaf gas exchange and the differential tolerance to soil flooding of L. corniculatus and L. glaber. Adult plants of these species, established independently in grasslands mesocosms, were subjected to 40 days of early spring flooding at a water depth of 6 cm. Both species presented constitutive aerenchyma tissue in the roots. Under flooding conditions, this parameter was 26.2% in L. glaber and 15.3% in L. corniculatus. In addition, flooded plants of L. glaber presented a leaf biomass 47.5% higher above water while L. corniculatus showed a leaf biomass 59.6% lower in the same layer, in comparison to control plants. Flooded plants of L. glaber maintained the stomatal conductance (g s ) and transpiration rate (E) for 25 days, although these parameters reduce slightly to 40À60% in comparison to controls after 40 days of flooding. In this species, a reduction in photosynthesis (A) in flooding conditions was detected only on the last day of measurement. In L. corniculatus, the same parameters (g s , E and A) were affected by flooding since day 18 of treatment, and values reached 25À40% in comparison to control plants after 40 days of flooding. Flooding did not affect above-ground biomass in L. glaber; while in L. corniculatus, above-ground biomass was 35% lower than in control plants. Our results confirmed that L. glaber is more able to cope with flooding stress than L. corniculatus, even in the presence of natural competitors. On the whole, this experiment provides information that can aid in the identification of anatomical and physiological parameters associated with flood-tolerance in this forage legume species, with economic potential for the agricultural areas subject to periodic flooding.
Leaf epidermal peels of Arabidopsis (Arabidopsis thaliana) mutants lacking either phototropins 1 and 2 (phot1 and phot2) or cryptochromes 1 and 2 (cry1 and cry2) exposed to a background of red light show severely impaired stomatal opening responses to blue light. Since phot and cry are UV-A/blue light photoreceptors, they may be involved in the perception of the blue light-specific signal that induces the aperture of the stomatal pores. In leaf epidermal peels, the blue light-specific effect saturates at low irradiances; therefore, it is considered to operate mainly under the low irradiance of dawn, dusk, or deep canopies. Conversely, we show that both phot1 phot2 and cry1 cry2 have reduced stomatal conductance, transpiration, and photosynthesis, particularly under the high irradiance of full sunlight at midday. These mutants show compromised responses of stomatal conductance to irradiance. However, the effects of phot and cry on photosynthesis were largely nonstomatic. While the stomatal conductance phenotype of phot1 phot2 was blue light specific, cry1 cry2 showed reduced stomatal conductance not only in response to blue light, but also in response to red light. The levels of abscisic acid were elevated in cry1 cry2. We conclude that considering their effects at high irradiances cry and phot are critical for the control of transpiration and photosynthesis rates in the field. The effects of cry on stomatal conductance are largely indirect and involve the control of abscisic acid levels.
The possibility that reduced photomorphogenic responses could increase field crop yield has been suggested often, but experimental support is still lacking. Here, we report that ectopic expression of the Arabidopsis PHYB (phytochrome B) gene, a photoreceptor involved in detecting red to far-red light ratio associated with plant density, can increase tuber yield in field-grown transgenic potato (Solanum tuberosum) crops. Surprisingly, this effect was larger at very high densities, despite the intense reduction in the red to far-red light ratios and the concomitant narrowed differences in active phytochrome B levels between wild type and transgenics at these densities. Increased PHYB expression not only altered the ability of plants to respond to light signals, but they also modified the light environment itself. This combination resulted in larger effects of enhanced PHYB expression on tuber number and crop photosynthesis at high planting densities. The PHYB transgenics showed higher maximum photosynthesis in leaves of all strata of the canopy, and this effect was largely due to increased leaf stomatal conductance. We propose that enhanced PHYB expression could be used in breeding programs to shift optimum planting densities to higher levels.The world population is predicted to reach 8 billion by 2025, and this, in combination with current trends in dietary composition, will result in a substantial increase of food demand. The ability to satisfy this demand will rely heavily on the genetic improvement of cultivated plants to increase yield potentials and/or yield stability (i.e. the constancy of performance; Khush, 2001). Although yield per unit area continues to increase for many crops, maximum yield is not showing such an obvious trend, and this has been interpreted as a result of yield potential approaching a ceiling, whereas yield stability continues to grow (Mann, 1999). Although some consider that this ceiling represents the highest physically attainable yield (Sinclair, 1993), others are more confident that incorporation of the current wealth of knowledge in plant biology will overcome the decreasing rate of increase in yield potential. However, genetic modifications that appear beneficial when analyzed at the cellular or plant level do not necessarily translate into higher yields in the field, either because of negative side effects or simply because they do not operate on the limitations under field conditions.As incident solar radiation penetrates the canopy of commercial crops, it suffers a gradual attenuation of the photosynthetic photon flux density (PPFD) and of the red (R) to far-red (FR) ratio (Holmes and Smith, 1977). Changes in the R to FR ratio perceived by phytochromes initiate a number of responses such as increased stem extension growth (Morgan and Smith, 1976), reduced branching (Deregibus et al., 1983), and accelerated leaf senescence (Rousseaux et al., 1999), which are collectively called the shade avoidance syndrome (Smith, 2000). In growing canopies, some of these changes anticipate com...
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