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1.  A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere 
Annals of Botany  2013;112(9):1911-1920.
Background and Aims
Ongoing global warming has been implicated in shifting phenological patterns such as the timing and duration of the growing season across a wide variety of ecosystems. Linear models are routinely used to extrapolate these observed shifts in phenology into the future and to estimate changes in associated ecosystem properties such as net primary productivity. Yet, in nature, linear relationships may be special cases. Biological processes frequently follow more complex, non-linear patterns according to limiting factors that generate shifts and discontinuities, or contain thresholds beyond which responses change abruptly. This study investigates to what extent cambium phenology is associated with xylem growth and differentiation across conifer species of the northern hemisphere.
Xylem cell production is compared with the periods of cambial activity and cell differentiation assessed on a weekly time scale on histological sections of cambium and wood tissue collected from the stems of nine species in Canada and Europe over 1–9 years per site from 1998 to 2011.
Key Results
The dynamics of xylogenesis were surprisingly homogeneous among conifer species, although dispersions from the average were obviously observed. Within the range analysed, the relationships between the phenological timings were linear, with several slopes showing values close to or not statistically different from 1. The relationships between the phenological timings and cell production were distinctly non-linear, and involved an exponential pattern
The trees adjust their phenological timings according to linear patterns. Thus, shifts of one phenological phase are associated with synchronous and comparable shifts of the successive phases. However, small increases in the duration of xylogenesis could correspond to a substantial increase in cell production. The findings suggest that the length of the growing season and the resulting amount of growth could respond differently to changes in environmental conditions.
PMCID: PMC3838565  PMID: 24201138
Cambium; cell differentiation; cell production; climate change; conifers; growth; meristem; phenology; productivity; secondary wall formation; xylogenesis
2.  Causes and correlations in cambium phenology: towards an integrated framework of xylogenesis 
Journal of Experimental Botany  2011;63(5):2117-2126.
Although habitually considered as a whole, xylogenesis is a complex process of division and maturation of a pool of cells where the relationship between the phenological phases generating such a growth pattern remains essentially unknown. This study investigated the causal relationships in cambium phenology of black spruce [Picea mariana (Mill.) BSP] monitored for 8 years on four sites of the boreal forest of Quebec, Canada. The dependency links connecting the timing of xylem cell differentiation and cell production were defined and the resulting causal model was analysed with d-sep tests and generalized mixed models with repeated measurements, and tested with Fisher’s C statistics to determine whether and how causality propagates through the measured variables. The higher correlations were observed between the dates of emergence of the first developing cells and between the ending of the differentiation phases, while the number of cells was significantly correlated with all phenological phases. The model with eight dependency links was statistically valid for explaining the causes and correlations between the dynamics of cambium phenology. Causal modelling suggested that the phenological phases involved in xylogenesis are closely interconnected by complex relationships of cause and effect, with the onset of cell differentiation being the main factor directly or indirectly triggering all successive phases of xylem maturation.
PMCID: PMC3295399  PMID: 22174441
Causal modelling; cell differentiation; cell production; d-sep test; Picea mariana; secondary wall formation; xylogenesis
3.  Dynamics of Depletion and Replenishment of Water Storage in Stem and Roots of Black Spruce Measured by Dendrometers 
In the short term, trees rely on the internal storage of water because it affects their ability to sustain photosynthesis and growth. However, water is not rapidly available for transpiration from all the compartments of the plant and the living tissues of the stem act as a buffer to preclude low water potentials during peaks of transpiration. In this paper, electronic dendrometers were used from mid-June to mid-September 2008 to compare the radius variations in stem and roots of black spruce [Picea mariana (Mill.) B.S.P.] in two sites of the boreal forest of Quebec, Canada, with different soil characteristics and water retention. The duration of the daily cycles was similar between sites and measurement heights but greater amplitudes of contraction and expansion were observed on the stem and in the site with the shallowest soil organic layer. The expansion phase had higher amplitudes and lasted longer than contraction. On average, the contraction phase occurred between 07:00 and 16:30 (legal time), while expansion lasted 14.5 h. The roots in the site with the deepest organic layer showed a wider variation in the onset of contraction, which could be as late as 13:00. The probability of observing the contraction phase depended on precipitation. With a precipitation <0.5 mm h−1, the bivariate posterior probabilities estimated >60% probability of observing contraction between 05:00 and 21:00, decreasing to 20% with precipitation >1.1 mm h−1. These findings demonstrated that the depth of the organic layer plays an important role in maintaining the internal water reserve of trees. The dynamics of water depletion and replenishment can modify the water potential of xylem and cell turgor during the enlargement phase, thus affecting radial growth. Changes in temperature and precipitation regime could influence the dynamics of internal water storage in trees growing on shallower and drier soils.
PMCID: PMC3355585  PMID: 22639583
boreal forest; climate change; stem cycle approach; plant–water relationships; soil depth; water reserves
4.  Cambial Activity and Intra-annual Xylem Formation in Roots and Stems of Abies balsamea and Picea mariana 
Annals of Botany  2008;102(5):667-674.
Background and Aims
Studies on xylogenesis focus essentially on the stem, whereas there is basically no information about the intra-annual growth of other parts of the tree. As roots strongly influence carbon allocation and tree development, knowledge of the dynamics of xylem production and maturation in roots at a short time scale is required for a better understanding of the phenomenon of tree growth. This study compared cambial activity and xylem formation in stem and roots in two conifers of the boreal forest in Canada.
Wood microcores were collected weekly in stem and roots of ten Abies balsamea and ten Picea mariana during the 2004–2006 growing seasons. Cross-sections were cut using a rotary microtome, stained with cresyl violet acetate and observed under visible and polarized light. The number of cells in the cambial zone and in differentiation, plus the number of mature cells, was counted along the developing xylem.
Key Results
Xylem formation lasted from the end of May to the end of September, with no difference between stem and roots in 2004–2005. On the contrary, in 2006 a 1-week earlier beginning of cell differentiation was observed in the stem, with cell wall thickening and lignification in roots ending up to 22 d later than in the stem. Cell production in the stem was concentrated early in the season, in June, while most cell divisions in roots occurred 1 month later.
The intra-annual dynamics of growth observed in stem and roots could be related to the different amount of cells produced by the cambium and the patterns of air and soil temperature occurring in spring.
PMCID: PMC2712372  PMID: 18708643
Abies balsamea; boreal forest; cambium; cell differentiation; cell wall thickening; lignification; Picea mariana; root; stem; xylem

Results 1-4 (4)