Aim To identify temperatures at which cell division and differentiation are active in order to verify the existence of a common critical temperature determining growth in conifers of cold climates. Location Ten European and Canadian sites at different latitudes and altitudes. Methods The periods of cambial activity and cell differentiation were assessed on a weekly time-scale on histological sections of cambium and wood tissue collected over 2 to 5 years per site from 1998 to 2005 from the stems of seven conifer species. All data were compared with daily air temperatures recorded from weather stations located close to the sites. Logistic regressions were used to calculate the probability of xylogenesis and of cambium being active at a given temperature. Results Xylogenesis lasted from May to October, with a growing period varying from 3 to 5 months depending on location and elevation. Despite the wide geographical range of the monitored sites, temperatures for onset and ending of xylogenesis converged towards narrow ranges with average values around 4-5, 8-9 and 13-14 degrees C for daily minimum, mean and maximum temperature, respectively. On the contrary, cell division in the cambium stopped in July-August, when temperatures were still high. Main conclusions Wood formation in conifers occurred when specific critical temperatures were reached. Although the timing and duration of xylogenesis varied among species, sites and years, the estimated temperatures were stable for all trees studied. These results provide biologically based evidence that temperature is a critical factor limiting production and differentiation of xylem cells in cold climates. Although daily temperatures below 4-5 degrees C are still favourable for photosynthesis, thermal conditions below these values could inhibit the allocation of assimilated carbon to structural investment, i.e. xylem growth
40The interaction between xylem phenology and climate assesses forest growth and productivity 41 and carbon storage across biomes under changing environmental conditions. We tested the annual temperature, from 83.7 days at -2 °C to 178.1 days at 12 °C, at a rate of 6.5 days °C -1 .
54April-May temperatures produced the best models predicting the dates of wood formation.
55Our findings demonstrated the uniformity of the process of wood formation and the 56 importance of the environmental conditions occurring at the time of growth resumption.
57Under warming scenarios, the period of wood formation might lengthen synchronously in the 58
The seasonal dynamic s of cambial activity, wood formation , occurrence of false rings (FR), and tangential bands of resin canals (RC) were investigated in Pinus halepensis from three Mediterranean dry and semi-arid ecosystems in Spain. We collected intact tissues of phloem, cambium, and outer xylem at monthly intervals throughout 2004 from each time six trees at the three sites. Cell divisions in the cambium in all trees started before our first sampling in mid-March and ceased between November and December. Cambial activity was characterized by two maxima; one in spring and another in autumn. Trees still grew in summer but at a very low rate. The first solitary RC were formed in May or June and tangential bands ofRC in June or July. In general , tangential bands ofRC were observed in wider growth rings . The formation of tangential bands of RC seems to be induced by drought in the second half of the growing season . FR were formed as a result of cambial reactivation in autumn and were observed in growth rings that contained more than 50 cells in a radial row.It appears that summer drought and early autumn precipitation play an important role in false-ring formation.
Heating and cooling can influence cambial activity and cell differentiation in Norway spruce. However, at the ultrastructural and topochemical levels, no changes were observed in the pattern of secondary cell-wall formation and lignification or in lignin structure, respectively.
Long-term variation in tree-ring widths and intra-annual dynamics of cambial activity and tree-ring formation in 2006 were studied in mature beech (Fagus sylvatica L.) trees at a typical forest site near Ljubljana (46°N, 14°40 0 E, 400 m a.s.l.) and related to leaf phenology and climate data. Tree-ring widths were nega-
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