Climate change is expected to be heterogeneous across the world, with high impacts on the Himalayan ecosystems. There is a need to precisely document cambial phenology and wood formation in these regions to better understand climate-growth relationships and how trees face a warming climate. This study describes the dynamics of cambial phenology in pindrow fir (Abies pindrow) along its altitudinal gradient in the Himalaya. The stages of xylem phenology, and the duration and rate of wood formation were assessed from anatomical observations during the growing season from samples collected weekly from three sites at various altitudes (2392–2965 m a.s.l.) over two years. There were significant differences in the duration and rate of cell formation along the altitudinal gradient, which decreased at increasing altitudes. The growing season duration decreased by 5.2 and 3.7 days every 100 m of increase in altitude in 2014 and 2015, respectively, while the rate of cell formation decreased from 0.38 and 0.44 cells /day to 0.29 and 0.34 cells/day in 2014 and 2015, respectively. Cell production decreased from 63.3 and 67.0 cells to 38.3 and 45.2 cells with a decrease of 4.3 and 3.8 cells per 100 m increase in altitude in 2014 and 2015, respectively. The higher precipitation in 2015 increased the growth rate and resulted in a higher xylem production. Our findings give new insights into the dynamics of cambial phenology and help in better understanding of the potential impacts of climate change on tree growth and forest productivity of Himalayan forests.
We monitored six healthy dominant trees and six girdled Scots pine trees for two successive growing seasons (2014 and 2015) to investigate the seasonal dynamics, cambial activity, and morphology of the new xylem and phloem cells formed under environmental stress when girdling was applied during the dormant period (15 January 2014). Microcore (1.8 mm) samples were collected weekly using a Trephor tool above and below the girdling area, and weather data were measured on site. Drought stress in combination with girdling reduced the total number of differentiation days cell formation. In 2014, no significant differences in tracheid dimensions were observed between the girdled area and the control trees, while in 2015, the control trees showed significantly smaller cell wall thickness and radial dimensions of the latewood tracheids (LW) compared to 2014 and girdled trees had no occurrence of LW. Under stressful heat waves and prolonged periods of no precipitation, the trees tended to reduce the number of tracheids that were formed and exhibited smaller radial dimensions (narrower tree rings) to increase their hydraulic efficiency. Trees responded to limited water availability by forming intra-annual density fluctuations (IADFs L) in the zone of the LW to overcome stressful conditions. Although xylem cell differentiation was affected by stressful conditions, no significant variability in phloem cell dimensions was observed. Thus, the phloem tissue was less sensitive to exogenous factors.
Although cell-anatomical variables are promising proxies reflecting seasonal as well as annual climate changes, their interdependencies are not yet fully understood. In the present study we assessed the changes in tree-ring width and various wood anatomical traits, including wall thickness, lumen diameter and tracheid diameter in the radial direction in saplings of Pinus sylvestris under six climatic conditions: 5°C warmer alone (ET) or combined with drought in June (ETJ) and in August (ETA) and CO2 enrichment alone (EC, 770 ppm) or combined with drought in June (ECJ) and in August (ECA). The experiments related to temperature conditions using 2-year saplings and CO2 conditions using 3-year saplings were completed in 2009 and 2010 in a greenhouse, respectively. Results showed that tree-ring width and tracheid diameter were not affected by any of the conditions applied, but the lumen diameter was larger and the wall thickness was thinner than those under control conditions. These reactions were verified under ETJ in the warming treatment and under all conditions under CO2 enrichment conditions. Our results indicated that drought counteracted the effects of elevated CO2 concentrations on wood anatomical properties, signifying complex interactions between the two major effects of climate change. Our comparison of wood parameters through experiments highlight the potential effect of climate change — increased drought stress due to higher temperatures and water shortage as well as elevated ambient CO2, on tracheid lumen diameter and wall thickness. Whereas the ring-width and tracheid diameter practically remained unaffected under the above-mentioned conditions.
Not only are vessel pits vital for the passage of sap into adjacent cells, but their anatomical morphology is also used as a tool to identify bamboo species. However, detailed studies comparing the pits’ structural parameters in culms of species with three rhizome types: sympodial bamboo, amphipodial bamboo, and monopodial bamboo, are lacking. Scanning electron microscopy (SEM) observations were conducted to obtain the qualitative and quantitative characteristics of vessel pits in sympodial, amphipodial and monopodial bamboos, from twelve bamboo species in eight genera. Sympodial bamboos possess small and ovoid bordered pits, whereas amphipodial bamboos contain an abundance of slit-like pits, with the greatest pit membrane length occurring in the vessel wall. Both minute and large pit sizes can be found in monopodial bamboos. This study identified the first compound pits ever to be found in a bamboo species and these were found to occur more frequently in amphipodial and monopodial bamboos than in sympodial bamboos. Using the distribution frequency of the pit chamber’s horizontal diameter, we were able to determine pit size as being either small, medium or large. The striking differences in the vessel pits’ qualitative and quantitative characteristics could be the result of different climate and environmental factors.
Recent studies on the seasonal dynamics of secondary tissue formation in Mediterranean trees have shown that xylogenesis depends on species and site conditions, but many questions still remain open. On the other side of the cambium, even less information is available about phloem structure and timing of its formation. We analysed intra-annual phloem variation in width and cell traits in the conducting, non-collapsed phloem (CPH) of Pinus pinea and Pinus halepensis at Mediterranean sites in southern Italy and Spain. In all investigated trees, it was possible to differentiate among the non-conducting, collapsed phloem (NCPH), and the CPH. CPH showed no evident annual growth layers; no differences in radial dimensions of early- and late phloem sieve cells, and no cyclic patterns of axial parenchyma distribution. Since it was not possible to study the seasonality of the phloem growth, we analysed the entire CPH. CPH width showed seasonal fluctuations and was generally the widest during the maximum cambial activity and narrowest during summer and winter. The radial size of newly formed sieve cells varied in relation to seasonal dynamics of cambial activity and fluctuations in local weather conditions. The number of axial parenchyma cells in CPH increased during the summer. The observed intra-annual variations in CPH width and structure seemed to be correlated with seasonal weather conditions in order to ensure a sufficient amount of conducting phloem tissue for translocation of photosynthates and signalling molecules to the actively growing tissues along the stem of a tree growing in the harsh Mediterranean conditions.
As the global climate warms, increased aridity is expected to become a major determinant of forest productivity and tree growth. In gymnosperms, wood density quantified at seasonal to annual scales can be related to changes in tracheid lumen size due to alterations in soil water availability. In this way, minimum wood density (MND) has been shown to respond negatively to early growing-season precipitation in several conifers because dry conditions reduce tracheid lumen size and consequently increase MND. We investigated if this relationship between spring precipitation and MND applies to four conifer species (Abies alba, Pinus sylvestris, Pinus nigra, Juniperus thurifera) in NE Spain from mesic (A. alba, P. sylvestris) to xeric (P. nigra, J. thurifera) conditions. We further assessed how climate, precipitation, and drought-affected tree-ring width (TRW) and MND at several time scales to test if water shortage in spring increases MND and decreases TRW over time and seasonally. Lastly, we quantified the post-drought MND recovery. We found the strongest negative correlations between MND and spring precipitation in P. nigra followed by J. thurifera. In these two species, the associations between MND and 9-month long droughts peaked in early spring (P. nigra, ; J. thurifera, ). Juniperus thurifera presented a better post-drought recovery (decrease in MND), followed by P. nigra and P. sylvestris. We conclude that MND is a reliable and accurate proxy of drought severity during spring in conifers subjected to seasonal water shortage. MND can be used as an early-warning indicator of short- and long-term changes in the responses of trees to water shortage.
This study assessed whether allometric scaling applied to pit sizes in stems of Douglas fir. Pit and pit aperture diameters were measured in xylem from stems of four plantation-grown Douglas fir (Pseudotsuga menziesii) trees from each of two different sites in Denmark. One site had fertile soil, the other had poor soil fertility. Three different heights were accessed for each tree, and 40 pits were measured per height. Results showed that pit diameter varied between 17 and 24 μm and decreased significantly with increasing height above ground. Representing the position in the tree as a power function of distance from the top of the tree (L0.2) rather than the height above ground improved the model fit for pit diameter as expressed by the R2 value. However, the pit diameter relationship was found to be significantly affected by site, suggesting that anatomic dimensions relate not only to tree size but also to growth conditions. This would imply that even though pit size supposedly has a strong biophysical determination, some xylem plasticity may still be induced by environmental factors.