Leaf gas exchange and plant water relations of three co-occurring evergreen Mediterranean shrubs species, Quercus ilex L. and Phillyrea latifolia L. (typical evergreen sclerophyllous shrubs) and Cistus incanus L. (a drought semi-deciduous shrub), were investigated in order to evaluate possible differences in their adaptive strategies, in particular with respect to drought stress. C. incanus showed the highest annual rate of net photosynthetic rate (PN) and stomatal conductance (gs) decreasing by 67 and 69 %, respectively, in summer. P. latifolia and Q. ilex showed lower annual maximum PN and gs, although PN was less lowered in summer (40 and 37 %, respectively). P. latifolia reached the lowest midday leaf water potential (Ψ1) during the drought period (-3.54±0.36 MPa), 11 % lower than in C. incanus and 19 % lower than in Q. ilex. Leaf relative water content (RWC) showed the same trend as Ψ1. C. incanus showed the lowest RWC values during the drought period (60 %) while they were never below 76 % in P. latifolia and Q. ilex; moreover C. incanus showed the lowest recovery of Ψ1 at sunset. Hence the studied species are well adapted to the prevailing environment in Mediterranean climate areas, but they show different adaptive strategies that may be useful for their co-occurrence in the same habitat. However, Q. ilex and P. latifolia by their water use strategy seem to be less sensitive to drought stress than C. incanus. and A. Bombelli, L. Gratani.
There are significant differences in leaf life-span among evergreen sclerophyllous species and drought semideciduous species growing in the Mediterranean maquis. Cistus incamus, which has a leaf life-span of four-eight months, was characterised by the highest net photosynthetic rates (PN), while Quercus ilex and Phillyrea latifolia, which maintain their leaves two-three and two-four years, respectively, had a lower PN. The longer leaf life-span of the two evergreen sclerophyllous species may be justified to cover the high production costs of leaf protective structures such as cuticle, hairs, and sclereids: cuticle and hairs screen radiation penetrating into the more sensitive tissues, and sclereids have a light-guiding function. Q. ilex and P. latifolia have the highest leaf mass/area ratio (LMA = 209 g m-2) and a mesophyll leaf density (2065 cells per mm2 of leaf cross section area) about two times higher than C. incanus. In the typical evergreen sclerophyllous species the steepest leaf inclination (α = 56°) reduces 42 % of radiation absorption, resulting in a reduced physiological stress at leaf level, particularly in summer. C. incanus, because of its low leaf life-span, requires a lower leaf investment in leaf protective structures. It exhibits a drastic reduction of winter leaves just before summer drought, replacing them with smaller folded leaves. The lower leaf inclination (α = 44°) and the lower LMA (119 g m-2) of C. incanus complement photosynthetic performance. Water use efficiency (WUE) showed the same trend in Q. ilex, P. latifolia, and C. incanus, decreasing 60 % from spring to summer, due to the combined effects of decreased CO2 uptake and increased transpirational water loss. and L. Gratani, A. Bombelli.
The morphological, anatomical and physiological variations of leaf traits were analysed during Quercus ilex L. leaf expansion. The leaf water content (LWC), leaf area relative growth rate (RGRl) and leaf dry mass relative growth rate (RGRm) were the highest (76±2 %, 0.413 cm2 cm-2 d-1, 0.709 mg mg-1 d-1, respectively) at the beginning of the leaf expansion process (7 days after bud break). Leaf expansion lasted 84±2 days when air temperature ranged from 13.3±0.8 to 27.6±0.9 °C. The net photosynthetic rate (PN), stomatal conductance (g s), and chlorophyll content per fresh mass (Chl) increased during leaf expansion, having the highest values [12.62±1.64 µmol (CO2) m-2 s-1, 0.090 mol (H2O) m-2 s-1, and 1.03±0.08 mg g-1, respectively] 56 days after bud break. Chl was directly correlated with leaf dry mass (DM) and P N. The thickness of palisade parenchyma contributed to the total leaf thickness (263.1±1.5 µm) by 47 %, spongy layer thickness 38 %, adaxial epidermis and cuticle thickness 9 %, and abaxial epidermis and cuticle thickness 6 %. Variation in leaf size during leaf expansion might be attributed to a combination of cells density and length, and it is confirmed by the significant (p<0.001) correlations among these traits. Q. ilex leaves reached 90 % of their definitive structure before the most severe drought period (beginning of June - end of August). The high leaf mass area (LMA, 15.1±0.6 mg cm-2) at full leaf expansion was indicative of compact leaves (2028±100 cells mm-2). Air temperature increasing might shorten the favourable period for leaf expansion, thus changing the final amount of biomass per unit leaf area of Q. ilex. and L. Gratani, A. Bonito.