SUMMARY
Article Details: Received: 2020-03-28 | Accepted: 2020-05-22 | Available online: 2020-09-30 https://doi.org/10.15414/afz.2020.23.03.125-138Leaves are part of the plant organs that are important to sustain its life. These organs are sensitive to climate changes and may present phenotypic plasticity in response to environmental conditions. However, affirmation of the leaves morphological plasticity and their regulation in different environments is still little studied up to date. In the present research, we evaluated performance of 20 different groups of Argania spinosa (L.) Skeels genotypes (half-sibling). Each group contains 3 half-sibs. Genotype × environment interactions (GxE) were evaluated as well, for shape and size leaves. To perform this, geometric morphometric principles were applied to analyze genotypes morphology in three locations (Central, North-Western and South-Western of Morocco). Univariate and multivariate analysis was used for data analysis. Results showed significant variation of symmetric and asymmetric components for genotypes, half-sibling and location with relatively high variation coefficient (ca 60%). Shape and size differences among genotypes, suggest that they were the main source in leaf morphology variation. Canonical Variate Analysis of leaf shapes reveals that the regions are clearly distinct from each other. For symmetric component analysis, Mahalanobis distances values among locations reached 35.53 between South-Western and North-Western locations, 21.88 North-Western and Central locations and 18.29 for South-Western and Central location. The differentiation between the groups using the Canonical Variations value showed a significant effect of the environment on the studied argan tree genotypes. Small leaves and narrow blades were observed in Central location compared to others. However, leaves originated from South-Western location had mainly an ovate shape. The same genotypes presented a high spectrum of shape variation varying from obovate to ovate in the other regions. This study highlights the strong correspondence between leaf morphology and genotype within different environments, and demonstrates that GxE interaction shave an impact to take into consideration in breeding programs.Keywords: adaptation, Argania spinosa, environment, genotype, geometric morphometrics, leaf morphologyReferences BENGOUGH, A.G. et al. (2011). Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits. Journal of Experimental Botany, 62(1), 59–68. https://doi.org/10.1093/jxb/erq350BLOOM, A. J. et al. (1985). Resource limitation in plants An economic analogy. Annual Review of Ecology, Evolution and Systematics, 16, 363–392. https://doi.org/10.1146/annurev.es.16.110185.002051BRUSCH, P. et al. (2003). Within and among tree variation in leaf morphology of Quercus petraea (Matt.) Liebl. Natural populations. Trees, 17, 164–172. https://doi.org/10.1007/s00468-002-0218-yBUNN, S.M. et al. (2004). Leaf-level productivity traits in Populus grown in short rotation coppice for biomass energy. Forestry, 77, 307-323. https://doi.org/10.1093/forestry/77.4.307CHITWOOD, D.H.et al. (2013). A quantitative genetic basis for leaf morphology in a set of precisely defined tomato introgression lines. Plant Cell, 25, 2465–2481. https://doi.org/10.1105/tpc.113.112391COLEMA, J. S. et al. (1994). Interpreting phenotypic variation in plants. Ecology and Evolution, (Personal Edition) 9, 187–191. https://doi.org/10.1016/0169-5347(94)90087-6CORNU, M. (1897). Note on the structure of Moroccan argan fruits (Argania syderoxylon (L.) Skeels). Bulletin de la Société Botanique de France, 44, 181–187. https://doi.org/10.1080/00378941.1897.10830758CRICK, J. C. and GRIME, J.P. (1987). Morphological plasticity and mineral nutrient capture in two herbaceous species of contrasted ecology. New Phytologist, 107, 403–414. https://doi.org/10.1111/j.1469-8137.1987.tb00192.xDONG, M. et al. (1996). Morphological responses to nutrient availability in four clonal herbs. Vegetatio, 123, 183–192. https://doi.org/10.1007/BF00118270DONOVAN, L.A. et al. (2011). The evolution of the world wide leaf economics spectrum. Trends in Ecology and Evolution, 26, 88–95. https://doi.org/10.1016/j.tree.2010.11.011DRYDEN, I.L. and MARDIA, K.V. (1998). Statistical Shape Analysis. Wiley: Chichester.DUDLEY, S.A. and SCHMITT, J. (1995). Genetic differentiation in morphological responses to simulated foliage shade between populations of Impatiens capensis from open and woodland sites. Functional Ecology, 9, 655-66. https://doi.org/10.2307/2390158EL ABOUDI, A. (1990). Typology of sub-Mediterranean arganeraies and ecophysiology of the argan tree (Argania spinosa (L.) Skeels) in Souss region (Morocco): PhD thesis. Grenoble: EsSciences in University Joseph Fourier, France.EL AICH, A. et al. (2007). Ingestive behaviour of grazing goats in the southwestern argan forest of Morocco. Small Ruminant Rasearch, 70(2–3), 248–256. https://doi.org/10.1016/j.smallrumres.2006.03.011GILES YOUNG I.A. et al. (2017). Analysing phenotypic variation in Eucalyptus pauciflora across an elevation gradient in the Australian Alps. Researching functional ecology in Kosciuszko National Park. Field Studies in Ecology, 1, 17–25. GRATANI, L. (2014). Plant phenotypic plasticity in response to environmental factors. Advances in Botany, 2014, ID 208747. https://doi.org/10.1155/2014/208747GUET, J. et al. (2015). Genetic variation for leaf morphology, leaf structure and leaf carbon isotope discrimination in European populations of black poplar (Populus nigra L.). Tree Physiology, 35, 850–863. https://doi.org/10.1093/treephys/tpv056HAJLAOUI, I. et al. (2007). Effets des basses températures et de la photopériode sur la croissance et le développement inflorescentiel du fraisier non remontant. Tropicultura, 25(2), 82–86.HENDRICKSON, L. et al. (2004). Low temperature effects on photosynthesis and growth of grapevine. Plant Cell Environment, 27, 795–809. https://doi.org/10.1111/j.1365-3040.2004.01184.xIBAÑEZ, C. et al. (2017). Ambient temperature and genotype differentially affect developmental and phenotypic plasticity in Arabidopsis thaliana. BMC Plant Biology, 17(1), 114. https://doi. org/10.1186/s12870-017-1068-5IWAIZUMI, R. et al. (1997). Correlation of length of terminalia of males and females among nine species of Bactrocera (Diptera: Tephritidae) and differences among sympatric species of B. dorsalis complex. Annals of the Entomological Society of America, 90, 664–666.JOEL, G. et al. (1994). Leaf morphology along environmental gradients in Hawaiian Metrosideros polymorpha. Biotropica, 26, 17–22. https://doi.org/10.2307/2389106KLINGENBERG, C. P. (2011). MorphoJ: an integrated software package for geometric morphometrics. Molecular Ecology Resources, 11(2), 353–357. https://doi.org/10.1111/j.1755-0998.2010.02924.xKLINGENBERG, C. P. et al. (2002). Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry. Evolution, 56, 1909–1920. https://doi.org/10.1111/j.0014-3820.2002.tb00117.xKLINGENBERG, C.P. and MCINTYRE, G.S. (1998). Geometric morphometrics of developmental instability: analyzing patterns of fluctuating asymmetry with Procrustes methods. Evolution, 52, 1363–1375. https://doi.org/10.1111/j.1558-5646.1998.tb02018.xKUNDU, S. K. and TIGERSTEDT, P. M. A. (1997). Geographical variation in seed and seedling traits of Neem (Azadirachta indica A. Juss.) among ten populations studied in growth chamber. Silvae Genetica, 46, 2–3.LANDE, R. (2009). Adaptation to an extraordinary environment by evolution of phenotypic plasticity and genetic assimilation. Journal of Evolutionary Biology, 22(7), 1435–1446. https://doi.org/10.1111/j.1420-9101.2009.01754.xLANGLADE, N.B. et al. (2005). Evolution through genetically controlled allometry space. Proceedings of the National Academy of Sciences of the U.S.A, 102, 10221–10226. https://doi. org/10.1073/pnas.0504210102LI, X. et al. (2015). Influences of environmental factors on leaf morphology of Chinese Jujubes. PLoS ONE,10, e0127825. https://doi.org/10.1371/journal.pone.0127825MARENCO, R. A. et al. (2006). Hydraulically based stomatal oscillations and stomatal patchiness in Gossypium hirsutum. Functional Plant Biology, 33(12), 1103–1113. https://doi.org/10.1071/FP06115M’HIRIT, O. et al. (1998). The argan tree, a fruit species, multipurpose forest Mardaga. Mardaga: Sprimont, Belgique.M‘HIRIT, O. (1989). The argan tree : a multipurpose forest fruit tree. Formation Forestière Continue, thème “l‘arganier“. Rabat: Station de Recherche Forestière, Morocco.PENNINGTON, T.D. (1991). The Genera of the Sapotaceae. Kew & London: Kew Publishing, Royal Botanic Gardens.PERROT, E. (1907). Shea, Argan and some other succulent Sapotaceae from Africa. Les végétaux utiles de l‘Afrique Tropicale Française, Fascicule II, 127–158.PIGLIUCCI, M. (2001). Phenotypic Plasticity: Beyond Nature and Nurture. Baltimore: Johns Hopkins University Press.POSSEN, B. J. et al. (2014). Variation in 13 leaf morphological and physiological traits within a silver birch (Betula pendula) stand and their relation to growth. Canadian Journal of Forest Research, 44, 657–665. https://doi.org/10.1139/cjfr-2013-0493PRENDERGAST, H.D.V. and WALKER, C.C. (1992). The argan: multipurpose tree of Morocco. The Kew Magazine, 9, 76–85.PYAKUREL, A. and WANG, J.R. (2013). Leaf morphological variation among paper birch (Betula papyrifera Marsh.) genotypes across Canada. Open Journal of Ecology, 3, 284. https://doi.org/10.4236/oje.2013.34033RIEUF, P. (1962). The Argan tree fungi. Les Cahiers de la Recherche Agronomique Rabat, 15, 1–25. ROHLF, F. J. (2000). On the use of shape spaces to compare morphometric methods. Hystrix, 11, 8–24.ROHLF, F.J. (2010). Tps Series. Department of Ecology and Evolution, State University of New York, Stony Brook, New York. Retrieved June 8, 2011 from http://life.bio.sunysb.edu/morph/ROYER, D. L. et al. (2008). Sensitivity of leaf size and shape to climate within Acer rubrum and Quercus kelloggii. New Phytologist, 179, 808–817. https://doi.org/10.1111/j.14698137.2008.02496.xROYER, D.L. et al. (2009). Phenotypic plasticity of leaf shape along a temperature gradient in Acer rubrum. PloS ONE, 10, e7653. https://doi.org/10.1371/journal.pone.0007653SAUVAGE, Ch. and VINDT, J. (1952). Flore du Maroc analytique, descriptive et illustrée. Spermatophytes, Fascicule I, Ericales, Primulales, Plombaginales, Ebénales et Contortales. Work of the Cherifien Scientific Institute. Rabat, Morocco: INRA (pp. 83–85).TIAN, F. et al. (2011). Genome-wide association study of leaf architecture in the maize nested association mapping population. Nature Genetics, 43, 159–162. https://doi.org/10.1038/ng.746TSUKAYA, H. (2005). Leaf shape: genetic controls and environmental factors. The International Journal of Developmental Biology, 49, 547–555. https://doi.org/10.1387/ijdb.041921htURIBE-SALAS, D. et al. (2008). Foliar morphological variation in the white oak Quercus rugosa Née (Fagaceae) along a latitudinal gradient in Mexico: Potential implications for management and conservation. Forest Ecology and Management, 256, 2121–2126. https://doi.org/10.1016/j.foreco.2008.08.002VAN KLEUNEN, M. and FISCHER, M. (2007). Progress in the detection of costs of phenotypic plasticity in plants. New Phytologist, 176(4), 727–730. https://doi.org/10.1111/j.1469-8137.2007.02296.xVISCOSI, V. and CARDINI, A. (2011). Leaf Morphology, Taxonomy and Geometric Morphometrics: A Simplified Protocol for Beginners. PLoS ONE, 10, e25630. https://doi.org/10.1371/journal.pone.0025630WARREN, C.R. et al. (2005). Does rainfall explain variation in leaf morphology and physiology among populations of red ironbark (Eucalyptus sideroxylon subsp. tricarpa) grown in a common garden? Tree Physiology, 25, 1369–1378. https://doi.org/10.1093/treephys/25.11.1369ZAHIDI, A. et al. (2013). Variability in leaf size and shape in three natural populations of Argania spinosa (L.) Skeels. International Journal of Current Research and Academic Review, 1, 13–25.