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Background and Aims The existence of sclerophyllous plants has been considered an adaptive strategy against different environmental stresses. Given that it literally means 'hard-leaved', it is essential to quantify the leaf mechanical properties to understand sclerophylly. However, the relative importance of each leaf trait for mechanical properties is not yet well established. Methods Genus Quercus is an excellent system to shed light on this because it minimizes phylogenetic variation while having a wide variation in sclerophylly. We measured leaf anatomical traits and cell wall composition, analysing their relationship with leaf mass per area and leaf mechanical properties in a set of 25 oak species. Key Results The upper epidermis outer wall makes a strong and direct contribution to the leaf mechanical strength. Moreover, cellulose plays a crucial role in increasing leaf strength and toughness. The principal component analysis plot based on leaf trait values clearly separates Quercus species into two groups corresponding to evergreen and deciduous species. Conclusions Sclerophyllous Quercus species are tougher and stronger owing to their thicker epidermis outer wall and/or higher cellulose concentration. Furthermore, section Ilex species share common traits, although they occupy different climates. In addition, evergreen species living in mediterranean-type climates share common leaf traits irrespective of their different phylogenetic origin. [ABSTRACT FROM AUTHOR]

Nowadays, evergreen sclerophyllous and winter-deciduous malacophyllous oaks with different paleogeographical origins coexist under Mediterranean-type climates, such as the mixed forests of the evergreen Quercus ilex subsp. rotundifolia Lam. and the winter-deciduous Quercus faginea Lam. Both Mediterranean oaks constitute two examples of contrasting leaf habit, so it could be expected that they would have different functional strategies to cope with summer drought. In this study, we analysed photosynthetic, photochemical and hydraulic traits of different organs for Q. faginea and Q. ilex subsp. rotundifolia under well-watered conditions and subjected to very severe drought. The coordinated response between photosynthetic and hydraulic traits explained the higher photosynthetic capacity of Q. faginea under well-watered conditions, which compensated its shorter leaf life span at the expense of higher water consumption. The progressive imposition of water stress evidenced that both types of Mediterranean oaks displayed different functional strategies to cope with water limitations. Specifically, the decrease in mesophyll conductance associated with edaphic drought seems to be the main factor explaining the differences found in the dynamics of net CO2 assimilation throughout the drought period. The sharp decline in photosynthetic traits of Q. faginea was coupled with a strong decrease in shoot hydraulic conductance in response to drought. This fact probably avoided extensive xylem embolism in the stems (i.e. 'vulnerability segmentation'), which enabled new leaf development after drought period in Q. faginea. By contrast, leaves of Q. ilex subsp. rotundifolia showed effective photoprotective mechanisms and high resistance to drought-induced cavitation, which would be related with the longer leaf life span of the evergreen Mediterranean oaks. The co-occurrence of both types of Mediterranean oaks could be related to edaphic conditions that ensure the maintenance of soil water potential above critical values for Q. faginea , which can be severely affected by soil degradation and climate change. [ABSTRACT FROM AUTHOR]

Leaf mass per area ( LMA) has been suggested to negatively affect the mesophyll conductance to CO2 ( gm), which is the most limiting factor for area-based photosynthesis ( AN) in many Mediterranean sclerophyll species. However, despite their high LMA, these species have similar AN to plants from other biomes. Variations in other leaf anatomical traits, such as mesophyll and chloroplast surface area exposed to intercellular air space ( Sm/ S and Sc/ S), may offset the restrictions imposed by high LMA in gm and AN in these species., Seven sclerophyllous Mediterranean oaks from Europe/North Africa and North America with contrasting LMA were compared in terms of morphological, anatomical and photosynthetic traits., Mediterranean oaks showed specific differences in AN that go beyond the common morphological leaf traits reported for these species (reduced leaf area and thick leaves). These variations resulted mainly from the differences in gm, the most limiting factor for carbon assimilation in these species., Species with higher AN showed increased Sc/ S, which implies increased gm without changes in stomatal conductance. The occurrence of this anatomical adaptation at the cell level allowed evergreen oaks to reach AN values comparable to congeneric deciduous species despite their higher LMA. [ABSTRACT FROM AUTHOR]

The accumulation of epicuticular waxes over stomata in Quercus coccifera L. contributes to a severe reduction in maximum stomatal conductance (gs,max) under Mediterranean (MED) conditions. However, this phenomenon was not observed in this species under temperate (TEM) conditions, which could lead to differences in the ability to assimilate CO2 between the sites. We hypothesise that the overall importance of such a reduction in gs,max on photosynthesis is modulated by other factors affecting carbon gain, mainly mesophyll conductance to CO2 (gm), through a plastic response to changes in environmental conditions (i.e., vapour pressure deficit, VPD, and mean daily quantum flux density, Qint). The results reveal that leaves grown at the TEM site did not show an increased ability for net CO2 assimilation (AN), mainly due to an equal gm at both sites. This fact is explained by a trade-off between an increased conductance of the gas phase (gias) and a reduced conductance of the liquid phase (gliq) at the TEM site compared with the MED site. In spite of the reduction in gs,max at the MED site, transpiration (E) did not diminish during midsummer to the levels of the TEM site due to a higher VPD found at the MED site, yielding a higher water use efficiency (AN/E) at the TEM site. Moreover, photosynthetic nitrogen use efficiency was also higher at the TEM site, indicating these leaves can reach similar values of AN with lower nitrogen investment that those at the MED site. These results suggest that Q. coccifera does not always use the main resources (water and nutrients) at leaf level as efficiently as possible. Moreover, the different patterns of resource use (in particular N), together with the functional plasticity, cannot overcome the morpho-functional constraints that limit photosynthetic activity, even under potentially favourable conditions. [ABSTRACT FROM AUTHOR]

ABSTRACT The adaptive benefit of stomatal crypts remains a matter of controversy. This work studies the effect on gas exchange of cuticular rims that overarch the stomatal pore in the Mediterranean species Quercus coccifera L. growing under Mediterranean (lower relative humidities and high summer temperatures) or oceanic conditions (higher daily relative humidities and mild temperatures). After microscopic assessment of the leaf surfaces and stomatal architecture, the impact of the cuticular 'cup' on gas exchange was evaluated by employing three-dimensional finite element models. Here, we provide evidence for a high plasticity of the Q. coccifera cuticular cup, with much larger vents under oceanic conditions compared to small vents under Mediterranean conditions. This structure adds a substantial fixed resistance thereby strongly decreasing gas exchange under Mediterranean conditions. The cuticular cup, which also increases leaf internal humidity, might buffer the rapid changes in vapour pressure deficit ( VPD) often observed under Mediterranean conditions. Since water loss of guard and adjacent epidermal cells regulates stomatal aperture, we suggest that this structure allows an efficient regulation of stomatal conductance and optimum use of resources under high VPD. This study provides evidence that plasticity of stomatal architecture can be an important structural component of hydraulic adaptation to different climate conditions. [ABSTRACT FROM AUTHOR]

Climate change models predict an increase in aridity, especially in the regions under Mediterranean-type climates such as the Mediterranean Basin. However, there is a lack of ecophysiological studies supporting the selection of the more drought-adapted ecotypes for reforestation programs. In this study, we analyzed the anatomical and functional adaptations of 18-month-old seedlings to drought on 16 Quercus suber L. populations from the southeastern rear edge of the species distribution in northern Tunisia growing in a common garden, in order to identify the most appropriate material to use in reforestations. The results evidenced that populations from more xeric habitats displayed the highest leaf dry mass per area (LMA) and lowest leaf area (LA) values, together with the largest increase in the bulk modulus of elasticity (Δε) in response to drought (i.e., elastic adjustment). On the other hand, some populations with intermediate values of aridity, LMA and LA displayed the sharpest increase in proline concentration (ΔPro), with a concomitant increase in osmotic potential at full turgor (Δπo) (i.e., osmotic adjustment). Therefore, two different strategies seem to drive the within-species variation of the studied Q. suber populations in response to water scarcity: (i) a water saver strategy for improving water stress tolerance through the maximization of the elastic adjustment; and (ii) a water spender strategy for maintaining water absorption and photosynthetic activity under moderate water stress through the maximization of the osmotic adjustment. We concluded that the higher elastic adjustment, together with reduced LA and increased LMA, implied a better performance under drought stress in the populations of Q. suber from more xeric habitats, which can be considered the most drought-adapted ecotypes and, consequently, the most appropriate for reforestation programs under an eventual increase in aridity. [ABSTRACT FROM AUTHOR]

Plant functioning and survival in drylands are affected by the combination of high solar radiation, high temperatures, low relative humidity, and the scarcity of available water. Many ecophysiological studies have dealt with the adaptation of plants to cope with these stresses in hot deserts, which are the territories that have better evoked the idea of a dryland. Nevertheless, drylands can also be found in some other areas of the Earth that are under the Mediterranean-type climates, which imposes a strong aridity during summer. In this review, plant species from hot deserts and Mediterranean-type climates serve as examples for describing and analyzing the different responses of trees and shrubs to aridity in drylands, with special emphasis on the structural and functional adaptations of plants to avoid the negative effects of high temperatures under drought conditions. First, we analyze the adaptations of plants to reduce the input of energy by diminishing the absorbed solar radiation through (i) modifications of leaf angle and (ii) changes in leaf optical properties. Afterwards, we analyze several strategies that enhance the ability for heat dissipation through (i) leaf size reduction and changes in leaf shape (e.g., through lobed leaves), and (ii) increased transpiration rates (i.e., water-spender strategy), with negative consequences in terms of photosynthetic capacity and water consumption, respectively. Finally, we also discuss the alternative strategy showed by water-saver plants, a common drought resistance strategy in hot and dry environments that reduces water consumption at the expense of diminishing the ability for leaf cooling. In conclusion, trees and shrubs living in drylands have developed effective functional adaptations to cope with the combination of high temperature and water scarcity, all of them with clear benefits for plant functioning and survival, but also with different costs concerning water use, carbon gain, and/or leaf cooling. [ABSTRACT FROM AUTHOR]

• Holm oak is an evergreen and sclerophyllous tree closely associated to Mediterranean-type climates. • However, the origin of this species can be found in pre-Mediterranean lineages, under tropical or subtropical climates. • To survive in Mediterranean-type climate implies to withstand dry and hot summer, cold winters or both. • The overall response of holm oak to stress factors (drought, high or low temperatures) suggests the existence of a marked within-species variability. • The current taxonomical consideration of holm oak as a complex of two different subspecies might go beyond morphological traits or geographical distribution. Q. ilex is a circum-Mediterranean species which must deal with two stressful periods throughout the year: summer and winter. However, the intensity of these abiotic stresses is very variable depending on the specific area of its distribution range. Besides, two subspecies are usually distinguished, even sometimes recognised as two different true species. Nevertheless, differences in the response to both summer and winter stresses among populations overstep the purely taxonomic question. Regarding to temperature, this species has shown a high resistance to both extremes. On the one hand, critical temperature at leaf level has been estimated in 50.2 °C for a provenance in the Iberian Peninsula (Soria, Spain), a temperature which even in the hottest areas of its distribution range is hardly ever reached. On the other hand, minimal temperature for the survival of the leaves has been estimated in −19.8 °C for Italian provenances and -26.6 °C for one of the coldest provenances (Soria, Spain). Similarly, these temperatures are quite rarely reachable in their respective provenances. Concerning to summer-drought, this may compromise xylem stability in severe cases of scarcity of water in the soil, inducing cavitation when water soil potential is too negative. In this sense, Q. ilex has demonstrated to have a high resistance to cavitation, represented by quite negative values of P 50 and P 88 when compared to other species. Besides, Iberian provenances (i.e. the most continental ones) have shown a higher resistance to cavitation than French or Italian ones. Likewise, holm oak may suffer cavitation during winter, but in this case by a completely different factor: freezing-thawing cycles. Winter-cavitation seems to be related to vessel size, with wider vessels being more vulnerable to cavitate than narrower ones. In spite of its paleotropical origin, Q. ilex has achieved to develop multiple adaptations and physiological strategies that has enabled it not only to withstand the hard climatic conditions imposed by Mediterranean climate, but to colonize and even dominate great part of the Mediterranean landscape. [ABSTRACT FROM AUTHOR]