Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 mol(CO₂) m^–2s^–1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 mol(CO₂) m^–2s^–1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 mol(CO2) m^–2s^–1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.This revised version was published online in March 2005 with corrections to the page numbers.     

Relationship between net photosynthesis and leaf respiration in Mediterranean evergreen species

The relationship between net photosynthetic (P _N) and leaf respiration (R) rates of Quercus ilex, Phillyrea latifolia, Myrtus communis, Arbutus unedo, and Cistus incanus was monitored in the period February 2006 to February 2007. The species investigated had low R and P _N during winter, increasing from March to May, when mean air temperature reached 19.2 °C. During the favourable period, C. incanus and A. unedo had a higher mean P _N (16.4±2.4 μmol m⁻² s⁻¹) than P. latifolia, Q. ilex, and M. communis (10.0±1.3 μmol m⁻² s⁻¹). The highest R (1.89±0.30 μmol m⁻² s⁻¹, mean of the species), associated to a significant P _N decrease (62 % of the maximum, mean value of the species), was measured in July (mean R/P _N ratio 0.447±0.091). Q₁₀, indicating the respiration sensitivity to short-term temperature increase, was in the range 1.49 to 2.21. Global change might modify R/P _N determining differences in dry matter accumulation among the species, and Q. ilex and P. latifolia might be the most favoured species by their ability to maintain sufficiently higher P _N and lower R during stress periods.     

Interspecific Differences of Leaf Gas Exchange and Water Relations of Three Evergreen Mediterranean Shrub Species

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 (P _N) and stomatal conductance (g _s) decreasing by 67 and 69 %, respectively, in summer. P. latifolia and Q. ilex showed lower annual maximum P _N and g _s, although P _N was less lowered in summer (40 and 37 %, respectively). P. latifolia reached the lowest midday leaf water potential (₁) 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 ₁. 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 ₁ 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.     

Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 μmol(CO₂) m^?2s^?1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 μmol(CO₂) m^?2s^?1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 μmol(CO2) m^?2s^?1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.     

Leaf Anatomy,Inclination, and Gas Exchange Relationships in Evergreen Sclerophqldous and Drought Semideciduous Shrub Species

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 (P _N), while Quercus ilex and Phillyrea latifolia, which maintain their leaves two-three and two-four years, respectively, had a lower P _N. 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 ( = 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 ( = 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.     

Long-time variations in leaf mass and area of Mediterranean evergreen broad-leaf and narrow-leaf maquis species

Morphological (dry mass, DM; surface area, LA; leaf mass per area, LMA), anatomical (leaf thickness, L), phenological (leaf life span, LL), and physiological (net photosynthetic rate, P _N) leaf traits of the evergreen species co-occurring in the Mediterranean maquis developing at Castelporziano (Rome) were tested. The correlation analysis indicated that LMA variation was tightly associated with LL variations: Cistus incanus and Arbutus unedo had a short LL (4±1, summer leaves, and 11±1 months, respectively) and low LMA (153±19 g m⁻²) values, Quercus ilex, Phillyrea latifolia, and Pistacia lentiscus high LMA (204±7 g m⁻²) and long LL (22±3 months), Erica arborea, Erica multiflora, and Rosmarinus officinalis a short LL (9±2 months) and an either high (213±29 g m⁻², R. officinalis and E. multiflora) or low (115±17 g m⁻², E. arborea) LMA. LMA values were significantly (p≤0.05) correlated with P _N (r≥0.68). In the tested species, LMA increased in response to the decrease of the total rainfall during the leaf expansion period. LMA variation was due to the unequal variation of DM and LA in the considered species. LMA is thus a good indicator of evergreen maquis species capability to respond to climate change, in particular to total rainfall decrease in the Mediterranean basin.     

Effects of winter cold stress on photosynthesis and photochemical efficiency of PSII of the Mediterranean <Emphasis Type="Italic">Cistus albidus</Emphasis> L. and <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

This study examined the photosynthetic and growth performances of potted plants of Cistus albidus L. and Quercus ilex L. submitted either to natural Mediterranean winter conditions or to mild greenhouse conditions. Plants grown outdoors exhibited lower light and CO₂-saturated CO₂ assimilation rates (A_sat) and apparent quantum yield (_i) than those indoors. Until mid-winter, C. albidus had higher A_sat than Q. ilex, but differences disappeared after a period of severe cold. Maximal photochemical efficiency of PSII (Fv/Fm) measured predawn was higher in C. albidus than in Q. ilex, and decreased throughout the season in outdoor plants. Fv/Fm also decreased at light saturation (A_sat) in both species. Fv/Fm was correlated with photosynthetic capacity and efficiency (quantum yield), but the resulting regression slopes were different between the two species. At the physiological level, C. albidus seemed to cope better with cold stress than Q. ilex. However, winter stress induced reduction of leaf absorptance, increased leaf mass per area, extensive leaf damage and high plant mortality in C. albidus. This suggests that the high performance of C. albidus leaves is not likely to be maintained for long periods of cold stress, and may therefore depend on continuous leaf replacement. Quercus ilex showed a conservative behaviour, with low net assimilation rates but greater leaf and plant survival than C. albidus.     

Drought tolerance in the Mediterranean species <Emphasis Type="Italic">Quercus coccifera,Quercus ilex,Pinus halepensis</Emphasis>, and <Emphasis Type="Italic">Juniperus phoenicea</Emphasis>

We investigated the strategies of four co-occurring evergreen woody species Quercus ilex, Quercus coccifera, Pinus halepensis, and Juniperus phoenicea to cope with Mediterranean field conditions. For that purpose, stem water potential, gas exchange, chlorophyll (Chl) fluorescence, and Chl and carotenoid (Car) contents were examined. We recognized two stress periods along the year, winter with low precipitation and low temperatures that led to chronic photoinhibition, and summer, when drought coincided with high radiation, leading to an increase of dynamic photoinhibition and a decrease of pigment content. Summer photoprotection was related to non-photochemical energy dissipation, electron flow to alternative sinks other than photosynthesis, decrease of Chl content, and proportional increase of Car content. Water potential of trees with deep vertical roots (Q. coccifera, Q. ilex, and P. halepensis) mainly depended on precipitation, whereas water potential of trees with shallow roots (J. phoenicea) depended not only on precipitation but also on ambient temperature.     

Variations in leaf respiration across different seasons for Mediterranean evergreen species

Leaf respiration (R _L) of evergreen species co-occurring in the Mediterranean maquis developing along the Latium coast was analyzed. The results on the whole showed that the considered evergreen species had the same R _L trend during the year, with the lowest rates [0.83 ± 0.43 μmol(CO₂) m^?2 s^?1, mean value of the considered species] in winter, in response to low air temperatures. Higher R _L were reached in spring [2.44 ± 1.00 μmol(CO₂) m^?2 s^?1, mean value] during the favorable period, and in summer [3.17 ± 0.89 μmol(CO₂) m^?2 s^?1] during drought. The results of the regression analysis showed that 42% of R _L variations depended on mean air temperature and 13% on total monthly rainfall. Among the considered species, C. incanus, was characterized by the highest R _L in drought [4.93 ± 0.27 μmol(CO₂) m^?2 s^?1], low leaf water potential at predawn (Ψ_pd= ?1.08 ± 0.18 MPa) and midday (Ψ_md = ?2.75 ± 0.11 MPa) and low relative water content at predawn (RWC_pd = 80.5 ± 3.4%) and midday (RWC_md = 67.1 ± 4.6%). Compared to C. incanus, the sclerophyllous species (Q. ilex, P. latifolia, P. lentiscus, A. unedo) and the liana (S. aspera), had lower R _L [2.72 ± 0.66 μmol(CO₂) m^?2 s^?1, mean value of the considered species], higher RWC_pd (91.8 ± 1.8%), RWC_md (82.4 ± 3.2%), Ψ_pd (?0.65 ± 0.28 MPa) and Ψ_md (?2.85 ± 1.20 MPa) in drought. The narrow-leaved species (E. multiflora, R. officinalis, and E. arborea) were in the middle. The coefficients, proportional to the respiration increase for each 10°C rise (Q₁₀), ranging from 1.49 (E. arborea) to 1.98 (A. unedo) were indicative of the different sensitivities of the considered species to air temperature variation.     

Dampening effects of long‐term experimental drought on growth and mortality rates of a Holm oak forest

Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long‐term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (?66.5%) and Q. ilex (?17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005–2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra‐ and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long‐term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long‐term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species‐specific responses to drought, what may eventually lead to a partial community shift favoring the more drought‐resistant species. However, we also report a dampening of the treatment effect on the two drought‐sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.     

Physiological,morphological and anatomical trait variations between winter and summer leaves of Cistus species

Morphological, anatomical and physiological summer and winter leaf traits of Cistus incanus subsp. incanus, C. salvifolius and C. monspeliensis growing at the Botanical garden of Rome were analyzed. With regard to differences between summer and winter leaves of the considered species, leaf thickness (L) was 21% higher in summer than in winter leaves (mean of the considered species) and this increase was mostly the result of the increased palisade parenchyma thickness over the spongy parenchyma one (24 and 16% higher in summer than in winter leaves, respectively). Leaf mass area (LMA) and leaf tissue density (LTD) were 38% and 17% higher in summer than in winter leaves, respectively (mean of the considered species). The photosynthetic rate (P_N), stomatal conductance (g_s) and chlorophyll content (Chl) of summer leaves were 54%, 17% and 14% lower, respectively, than in winter leaves. C. monspeliensis summer leaves had the highest LMA, LTD, adaxial cuticle thickness (14.6 ± 1.8 mg cm⁻², 1091 ± 94 mg cm⁻³, and 5.8 ± 1.7 μm, respectively) and the lowest mesophyll intercellular spaces (f_ias 38 ± 3%). Moreover, C. monspeliensis had the highest P_N in summer (2.6 ± 0.1 μmol m⁻² s⁻¹) and C. incanus the highest P_N and WUE (84% and 59% higher than the other species) in the favorable period, associated to a higher f_ias (42 ± 2%). C. salvifolius had the highest P_N (54% higher than the other species) in winter. The plasticity index could allow a better interpretation of the habitat preference of the considered species. The physiological plasticity (PI_p = 0.39, mean value of the considered species) was higher than the morphological (PI_m = 0.22, mean value) and anatomical (PI_a = 0.13, mean value) plasticity. Moreover, among the considered species, C. salvifolius and C. incanus are characterized by a larger PI_a (0.14, mean value) which seems to be correlated with their wider ecological distribution and the more favorable conditions of the environments where they naturally occur. The highest PI_m (0.29) of C. monspeliensis indicates that it can play a high adaptive role in highly stressed environments, like fire degraded Mediterranean areas in which it occurs.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

Water relations,gas exchange,and growth of resprouts and mature plant shoots of Arbutus unedo L. and Quercus ilex L.

Resprout and mature plant shoot growth, leaf water status and gas exchange behavior, tissue nutrient content, flowering, and production were studied for co-occurring shallow-rooted (Arbutus unedo L.) and deeprooted (Quercus ilex L.) Mediterranean tree species at the Collserola Natural Park in Northeast Spain Resprouts showed higher growth rates than mature plant shoots. During fall, no differences in eco-physiological performance of leaves were found, but mobilization of carbohydrates from burls strongly stimulated growth of fall resprouts compared to spring resprouts, despite low exposed leaf area of the fall shoots. During summer drought, resprouts exhibited improved water status and carbon fixation compared to mature plant shoots. Shoot growth of Q. ilex was apparently extended due to deep rooting so that initial slower growth during spring and early summer as compared to A. unedo was compensated. Tissue nutrient contents varied only slightly and are postulated to be of minor importance in controlling rate of shoot growth, perhaps due to the relatively fertile soil of the site. Fall flowering appeared to inhibit fall shoot growth in A. unedo, but did not occur in Q. ilex. The results demonstrate that comparative examinations utilizing vegetation elements with differing morphological and physiological adaptations can be used to analyze relatively complex phenomena related to resprouting behavior. The studies provide an important multi-dimensional background framework for further studies of resprouting in the European Mediterranean region.     

Leaf morphology and leaf chemical composition in three Quercus (Fagaceae) species along a rainfall gradient in NE Spain

 Leaf features were examined in three Quercus species (Q. coccifera, Q. ilex and Q. faginea) along a steep rainfall gradient in NE Spain. The analyzed leaf traits were area, thickness, density, specific mass, leaf concentration of nitrogen, phosphorous, lignin, cellulose and hemicellulose, both on a dry weight basis (N_w, P_w, L_w, C_w, H_w) and on an area basis (N_a, P_a, L_a, C_a, H_a). These traits were regressed against annual precipitation and correlated with each other, revealing different response patterns in the three species. Q. faginea, a deciduous tree, did not show any significant correlation with rainfall. In Q. coccifera, an evergreen shrub, N_w, N_a, L_w, L_a and C_a increased with higher annual rainfall, while H_w decreased. In Q. ilex, an evergreen tree, leaf area, P_w and L_w increased with precipitation, whereas specific leaf mass, thickness and H_a showed the reverse response. Correlations between the leaf features revealed that specific mass variation in Q. faginea and Q. coccifera could be explained by changes in leaf density, while in Q. ilex specific leaf mass was correlated with thickness. Specific leaf mass in the three species appeared positively correlated with all the chemical components on a leaf area basis except with lignin in Q. ilex and with P in Q. ilex and Q. faginea. In these two tree species P_w showed a negative correlation with specific leaf mass. It is suggested that each species has a different mechanism to cope with water shortage which is to a great extent related to its structure as a whole, and to its habit. Received: 18 December 1995 / Accepted: 8 March 1996     

Leaf traits variation during leaf expansion in <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

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 (RGR_l) and leaf dry mass relative growth rate (RGR_m) 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 (P _N), 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.     

Seasonal changes in the content of photosynthetic pigments in perennial grasses of cryolithic zone

Seasonal changes in the content of photosynthetic pigments and their ratio were studied in perennial gramineous (Psathyrostachys juncea Tzvel., Elymus sibiricus L., Bromopsis inermis Leys) and equisetaceous (Equisetum variegatum Schleich. ex Web., E. scirpoides Michx.) plants grown under conditions of cryolithic zone in Central and North-East Yakutia (the Pole of Cold). Simultaneously with growth in chlorophyll (Chl) content in all the studied species of perennial grasses of Central Yakutia, a considerable increase in the content of carotenoids (Car) of the xanthophyll cycle (XC) pigments was observed with the beginning of autumn temperature fall. Similar dependence was found in horsetails of the Pole of Cold region: the content of Car was increased by 1.2–3.0 times comparing to the summer values, whereas at the same time the concentration of oxygen-containing forms increased and the ratio of Chl to xanthophylls and Car decreased. In general, horsetails had typically a lower content of Chl and Car, which was determined not only by extremely harsh climatic conditions of the region but also by specific characteristics of Equisetaceae plants. Closer to winter time, the formation of secondary Car, rhodoxanthin, was also observed in the studied horsetail species. In this case, rhodoxanthin presumably plays more important antioxidant role than XC, which can not operate at constantly low temperatures. The obtained data suggest that one of the characteristics of cold hardening of perennial grasses under conditions of cryolithic zone of Yakutia is, presumably, the high level of primary, as well as secondary, Car with most profound antioxidant properties.     

Chloroplastidic pigments,gas exchange,and carbohydrates changes during <Emphasis Type="Italic">Carapa guianensis</Emphasis> leaflet expansion

Changes in chloroplastidic pigments, gas exchange and carbohydrate concentrations were assessed during the rapid initial expansion of C. guianensis leaflet. Leaves at metaphyll stage were tagged and assessments were carried out 14, 17, 20, 23, 27, and 31 days later. Pigments synthesis, distribution and accumulation were uniform among leaflet sections (basal, median and apical). Chlorophyll (Chl) a, Chl b, Chl (a+b), and total carotenoids (Car) concentrations were significantly increased after 27 days from metaphyll, and the most expressive increases were parallel to lower specific leaflet area. Chl a/b was lower on day 14 and it was increased on subsequent days. Negative net photosynthesis rate (P _N), and the lowest stomatal conductance (g _s) and transpiration (E) were registered on day 14, following significant increases on subsequent days. The Chl (a+b) and Chl a effects on P _N were more expressive until day 20. Intercellular to ambient CO₂ concentration ratio (C _i/C _a) was higher on day 14 and lower on subsequent days, and no stomatal limitation to CO₂ influx inside leaflets was observed. Leaflet temperature was almost constant (ca. 35°C) during leaflet development. Sucrose and starch concentrations were increased in parallel to increases in P _N. Altogether, these results highlight the main physiological changes during C. guianensis leaflet expansion and they should be considered in future experiments focusing on factors affecting P _N in this species.     

Acclimation of winter wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis>, cv. Yangmai 13) to low levels of solar irradiance

Winter wheat is a grass species widely planted in northern and central China, where the increase of aerosols, air pollutants and population density are causing significant reduction in solar irradiance. In order to investigate the adaptation of winter wheat (Triticum aestivum L., cv. Yangmai 13) to low irradiance conditions occurring in the downstream plain of the Yangtze River (China), plants were subjected to four solar irradiance treatments (100%, 60%, 40%, and 20% of environmental incident solar irradiance). Significant increases in chlorophyll (Chl) and xanthophyll (Xan) pigments, and decreases in Chl a/b and Xan/Chl ratios were observed in plants under low light. Light-response curves showed higher net photosynthetic rates (P _N) in fully irradiated plants, that also showed a higher light-compensation point. Shaded plants maintained high values of minimal fluorescence of dark-adapted state (F_o) and maximum quantum efficiency of PSII photochemistry (F_v/F_m) that assess a lower degree of photoinhibition under low light. Reduced irradiance caused decreases in effective quantum yield of PSII photochemistry (Φ_PSII), electron transport rate (ETR), and nonphotochemical quenching coefficient (q_N), and the promotion of excitation pressure of PSII (1 − q_P). The activities of the antioxidant enzymes superoxide dismutase and peroxidase were high under reduced light whereas no light-dependent changes in catalase activity were observed. Thiobarbituric acid reactive species content and electrolyte leakage decreased under shaded plants that showed a lower photooxidative damage. The results suggest that winter wheat cv. Yangmai 13 is able to maintain a high photosynthetic efficiency under reduced solar irradiance and acclimates well to shading tolerance. The photosynthetic and antioxidant responses of winter wheat to low light levels could be important for winter wheat cultivation and productivity.     

Photosynthetic Characteristics of Sun and Shade Leaves in the Canopy of Arbutus unedo L. Trees Exposed to In Situ Long-Term Elevated CO2

Photosynthetic parameters were studied in Arbutus unedo L. trees growing at either ambient (AC) or elevated EC (mean 465 μmol mol^-1) CO₂ concentration near a natural CO₂ vent in Orciatico, Italy Diurnal courses of net photosynthetic rate (P _N), ratio of variable to maximum chlorophyll fluorescence (F_v/F_m), and quantum yield of electron transport through photosystem 2 (Φ₂) were measured on sun and shade leaves. The contents of N, C, Ca, K, P, and chlorophyll (Chl) and specific leaf area (SLA) in these leaf categories were also determined. A morning peak and midday depression of P _N were found for both AC and EC sun leaves. Long-term EC caused little or no down-acclimation of P _N in sum leaves. The estimate of total daily CO₂ uptake was lower in AC leaves than in EC leaves. In shade leaves, it reached up to 70 % of the value of sun leaves. The F_v/F_m ratio showed decreasing trend in the morning, reached a minimum at midday (90 % of dawn value), and then increased in the afternoon. The EC had no effect on F_v/F_m either in sun or shade leaves. Plants grown near the CO₂ spring had lower Chl content, higher SLA, and higher Ca and K contents than plants grown under AC. This revised version was published online in June 2006 with corrections to the Cover Date.     

Photosynthetic activity and water use efficiency of dune species: the influence of air temperature on functioning

Differences in leaf traits among the dune species developing along the Latium coast were analysed. Cakile maritima Scop. subsp. maritima, Elymus farctus (Viv.) Runemark ex Melderis subsp. farctus, Ammophila arenaria (L.) Link subsp. australis (Mabille) Lainz, Ononis variegata L., Pancratium maritimum L., Eryngium maritimum L., and Anthemis maritima L. were considered. The considered species showed a similar net photosynthetic rate (P _N) and chlorophyll content (Chl) during the year, with a peak from the end of April to the middle of May [13.0±3.6 μmol (CO₂) m⁻² s⁻¹ and 0.63±0.21 mg g⁻¹, respectively, mean values of the considered species], favoured by air temperature in the range 13.3–17.5°C, and 6% of soil water availability. In June–July, the increase of air temperature (T_max = 28.4°C), associated with a lower water availability (42 mm, total rainfall of the period) and a 1% of soil water availability determined a significant decrease of P _N (59%, mean of the considered species) and Chl (38%), and an increase of the carotenoid (Car)/Chl ratio (59%). The significant correlation between P _N and stomatal conductance (g _s) (p<0.05) explained 67% of P _N variations. Moreover, the correlation between P _N and leaf temperature (T _l) underlined that the favourable T _l enabling 90–100% of the highest P _N for the considered species was within the range 23.4 to 26.6°C. P _N decreased below half of its maximum value when T _l was over 35.8 and 37.4°C for E. farctus subsp. farctus and A. arenaria subsp. australis, respectively and over 32.2°C for the other considered species (mean value). Leaf mass area (LMA) varied from 6.8 ± 0.7 mg cm⁻² (O. variegata) to 30.6 ± 1.6 mg cm⁻² (A. arenaria). PCA (principal component analysis) carried out using the considered morphological and physiological leaf traits underlined that the co-occurring species were characterised by different adaptive strategies: E. farctus and A. arenaria photosynthesized for a long period also when air temperature was over 35.8 and 37.4°C, respectively, because of their lower transpiration rates [E, 1.4 ± 0.1 mmol (H₂O) m⁻² s⁻¹], which seemed to be controlled by the highest LMA. On the contrary, A. maritima and C. maritima subsp. maritima had a higher P _N (on an average 52% higher than the others) in the favourable period, allowed by the highest succulence index (SI, 85.7 ± 9 mg cm⁻²) and the lower LMA. The results allowed us to hypothesize that A. arenaria and E. farctus might be at a competitive advantage relative to the other considered species with respect to the increase of air temperature, by their ability to photosynthesize at sufficient rates also during summer.