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Dimensions of anatomical and morphological attributes of leaves can influence physiological response to changes in environment over time. Linking structural attributes of leaves to crown position and tree size within naturally developing cohorts of trees can provide a clearer understanding of changes in crown morphology for a species. This study examined leaf anatomy and morphology of Betula papyrifera Marsh. growing in two stages of stand development of interior cedar-hemlock forest, northern British Columbia. Anatomical and morphological measurements of leaves located at six different positions within the crown were made on trees selected from stands that originated 15 years (small tree size) and 145 years (large tree size) after catastrophic fire. Leaf area and mass were measured in the field. Microscopic measures were made in the laboratory of thicknesses of leaves, cuticle, upper and lower epidermis, palisade, and spongy mesophyll. Stomatal density and stomatal aperture lengths were also determined. Leaf anatomy and morphology varied significantly with position in the crown and among size-classes of trees. Changes in leaf anatomy observed among positions within the crown reflect the changing availability of light and moisture experienced during crown development. For both size-classes of tree the largest anatomical dimensions of leaves were at the outer and uppermost parts of the crown, whilst the smallest were at the lower and innermost parts. Foliage of large trees (145 years) exhibit leaf attributes characteristic of the sun-shade dichotomy reported in the literature, but this was not shown for foliage of young saplings (15 years). For the small trees (15 years) the largest leaves were located at the top of the crown while the smallest were located at the bottom. This has been reported for many tropical pioneers but is in contrast to the usual sun-shade dichotomy of temperate pioneers. The large trees (145 years) followed the typical pattern with the smallest leaves at the top of the crown. Information from this study contributes to our understanding of foliar development of tree crowns by demonstrating how leaf structure changes with crown position and tree size under stand competition.Résumé : Les dimensions des caractères anatomiques et morphologiques des feuilles peuvent influencer les réactions physiologiques aux changements du milieu dans le temps. Le fait de relier les caractères structuraux des feuilles avec leur position dans la couronne et la dimension de l'arbre, parmi un ensemble d'arbres poussant naturellement, peut conduire à une meilleure compréhension des changements morphologiques au niveau de la couronne, pour une espèce donnée. Dans cette étude, les auteurs examinent l'anatomie et la morphologie foliaire du Betula papyrifera Marsh. poussant à deux étapes du développement d'un peuplement, à l'intérieur de la forêt de cèdres et pruches, au nord de la Colombie Britannique. Les mesures anatomiques et morphologiques localisées à six positions différentes à l'intérieur de la couronne ont été ...

This study examined the foliar response of putative hybrid Engelmann × white × Sitka spruce (Picea engelmannii Parry × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carr) needles in relation to crown position and across three stages of development (15, 55, and 145 years). We focused on the morphological and anatomical response, and used physiological measures (photosynthesis and stomatal conductance) to emphasize the important relationship between structure and function. We found that needles from the upper outer crown position were adaptated to reduce stress from evapotranspiration. Trees from the 15- and 55-year-old stands generally responded the most, and trees from the 145-year-old stand responded the least. As they mature, these spruce trees may have reduced ability to adapt to their environment. Some of our results contradict what the literature considers "typical" for sun-shade dimorphism in temperate forests. For example, in all stands, sun needles were broader than shade needles and, in the two younger stands, sun needles were larger in area, not smaller, than shade needles. Also, in the youngest stand, stomatal pores were longer on sun needles than on shade needles. There were no definite patterns in stomatal spacing with regard to crown position. Our results are indicative of the strategies adopted to increase competitiveness in a resource-limited environment. We suggest that, in the 15-year-old stand, neither water nor light is limiting; in the 55-year-old stand, both water and light are highly limiting; and in the 145-year-old stand, water is most limiting.Key words: drought, foliar plasticity, needle anatomy, photosynthesis, Picea, stand development, sun-shade.

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