Seasonal physiological responses of Argania spinosa tree from Mediterranean to semi-arid climate

Argania spinosa (the argan tree) is a slow-growing tree endemic of Morocco, growing on semi-arid areas where no other tree species can live. With the aim of predicting temporal changes in A. spinosa woodlands under a probable increase in aridity, we set off to investigate these questions: how do A. spinosa physiological attributes respond to variations in climatic conditions and seasonality, and which is the set of attributes that most affects tree response to environmental conditions? In three study sites, Beni Snassen (North), High-Atlas (Mountain) and Admine Forest in Agadir (Coastal), gas exchange measurements, photochemical efficiency, leaf water potential and different leaf attributes were monitored in February, July and November of 2006. The Mountain site presents the most continental climate. Trees in this site were the most stressed in summer, having the lowest midday leaf water potential values, photochemical efficiency and assimilation rates. We found a Ψ_md threshold around -4 MPa, below which stomatal conductance responds linearly to Ψ_md. Plants from the North area never reached this threshold during the study period. Although leaf pigments presented a clear seasonal pattern, leaves from Coastal trees exhibit the highest content for each season. The three study sites were separated by two discriminate functions obtained by canonical discriminant analysis. In summer, the Mountain population is separated from the other sites mainly by assimilation rate and F_v/F_m, while in winter transpiration rates and chlorophyll content are the main discriminant variables. Our study shows that A. spinosa trees adjust their physiological status and leaf attributes to environmental conditions allowing plants to thrive under a dry climate. Under a scenario of global change, the distribution of the argan tree likely shifts to milder areas.     

Physiological,morphological and allocation plasticity of a semi-deciduous shrub

The main objective of this study was to look into the phenotypic plasticity of the semi-deciduous Mediterranean shrub, Halimium halimifolium. We studied morphological, allocation and physiological traits to determine which characters were more plastic and contribute in a greater extent to the acclimation ability of the species.We present a phenotypic plasticity index for morphological, physiological and allocation traits, which we have applied in the most contrasted plant communities where the species grows naturally. Data published by Díaz Barradas, M.C., García Novo, F. [1987. The vertical structure of Mediterranean scrub in Doñana National Park (SW Spain). Folia Geobotanica Phytotaxonomica 22, 415–433; 1988. Modificación y extinción de la luz a través de la copa en cuatro especies de matorral en el Parque Nacional de Doñana. Monografias Instituto Pirenaico de Ecologia 4, 503–516; 1990. Seasonal changes in canopy structure in two mediterranean dune shrubs. Journal of Vegetation Science 1, 31–40.], Díaz Barradas, M.C., Zunzunegui, M., García Novo, F. [1999a. Autoecological traits of Halimium halimifolium in contrasted habitats under Mediterranean type climate. Folia Geobotanica 34, 189–208.] and Zunzunegui et al. [Zunzunegui, M., Díaz Barradas, M.C., García Novo, F. 1997. Autoecological notes of Halimium halimifolium. Lagascalia 19, 725–736. Sevilla, Spain; Zunzunegui, M., Díaz Barradas, M.C., Fernández Baco, L., García Novo, F. 1999. Seasonal changes in photochemical efficiency in leaves of Halimium halimifolium a Mediterranean semideciduous shrub. Photosynthetica 36, 17–31; Zunzunegui, M., Díaz Barradas, M.C., García Novo, F. 2000. Different phenotypic response of Halimium halimifolium in relation to groundwater availability. Plant Ecology 148, 165–174; Zunzunegui, M., Díaz Barradas, M.C., Aguilar, F., Ain-Lhout, F., Clavijo, A., García Novo, F. 2002. Growth response of Halimium halimifolium at four sites with different soil water availability regimes in two contrasted hydrological cycles. Plant and Soil 247, 271–28.] have been re-appraised and combined with original data.Phenotypic plasticity index showed that the highest plasticity of the species was physiological, and especially for the traits related with water control. This high physiological plasticity appears mainly between seasons (September–November) rather than between sites. Plasticity in allocation traits showed intermediate values between physiological and morphological traits.     

Seasonal Differences in Photochemical Efficiency and Chlorophyll and Carotenoid Contents in Six Mediterranean Shrub Species under Field Conditions

The effects of summer and winter stress on the chlorophyll and carotenoid contents and photosystem 2 efficiency were examined in six Mediterranean scrub species. These six species belong to two different plant functional types: drought semi-deciduous (Halimium halimifolium L., Rosmarinus officinalis L., Erica scoparia L.) and evergreen sclerophylls (Juniperus phoenicea L., Pistacia lentiscus L., Myrtus communis L.). Two sites with different water availability were chosen. In the xerophytic site, despite they belong to two different functional types, R. officinalis and J. phoenicea showed a similar response. These were the most affected species in summer. H. halimifolium showed optimal values of F_v/F_m and non-significant seasonal changes in xanthophyll content. In the mesic site, E. scoparia and M. communis were apparently the most affected species by winter climatic conditions. P. lentiscus presented a pattern similar to H. halimifolium, except for elevated F₀ values. In all the studied species, lutein plus zeaxanthin content was negatively correlated with F_v/F_m in summer and with leaf water potential, thus indicating that the thermal dissipation of energy was a general pattern for all species. Under stress, plant response is more species-specific than dependent on its functional type.     

Seasonal physiological plasticity and recovery capacity after summer stress in Mediterranean scrub communities

Under natural conditions the overlapping of multiple stressors may initiate coordinated ecophysiological responses in Mediterranean species. Seasonal plasticity may enable plants to better cope with adverse environmental conditions and/or resource variability. In this article, we study the seasonal responses of 12 woody species in two sites of differing water availability, in a Mediterranean-type climate. Plants were measured for water potential, photochemical efficiency, photosynthetic pigments and leaf proline content throughout the year. The results revealed that species presented different ecophysiological strategies, even when sharing the same area. In the xerophytic site, some species suffered severe water stress (−12 MPa and F _v/F _m lower than 0.3), while others exhibited optimal values of F _v/F _m and only moderate water stress. All the plants recovered after the first autumn rains. In the hygrophytic site, some sclerophyll species did not exhibit signs of water stress, but did suffer photoinhibition in clear winter days. A plasticity index was calculated to provide an integrated value of species plasticity. In summer, plasticity was higher in the xerophytic site, while in winter it was higher in the hygrophytic site. Ordination analysis of the physiological traits supports the traditional gradient of Mediterranean strategies from drought semideciduous to evergreen sclerophyll species, although spiny legume species formed an independent functional group. The functional responses of species clearly differ among plant communities according to the prevailing site stressors, but no unique pattern emerges. Species combine traits in broader strategies according to previous evolutionary story exhibiting a certain amount of trade among traits, each contributing to alleviate a part of the plant stress.     

Comparison of proline accumulation in two mediterranean shrubs subjected to natural and experimental water deficit

The effect of water stress on proline accumulation was tested in two contrasted species of Mediterranean scrub: Halimium halimifolium (L.) Willk and Pistacia lentiscus L. Leaf water potential, stomatal resistance and proline content have been measured both in experimental and in natural water stress conditions. Both species accumulated proline in their leaves when leaf water potential dropped below a threshold value of –3.0 MPa, under natural as well as under experimental conditions. In the field, however, a time-lag between decrease of leaf water potential and proline accumulation could be observed. In Halimium halimifolium, proline accumulation appeared to be associated with severe stress conditions as most plants with high proline contents suffered irreversible wilting, especially in the greenhouse. P. lentiscus showed a different pattern, accumulating proline at two different times of the year, as a response to cold or to drought. The results of our study indicated that the role of proline in this species, rather than an osmotic agent, seems to be more related to a protective action in cases of severe stress conditions.