Xylem and phloem phenology in co-occurring conifers exposed to drought.

KEY MESSAGE: Variability in xylem and phloem phenology among years and species is caused by contrasting temperatures prevailing at the start of the growing season and species-specific sensitivity to drought. ABSTRACT: The focus of this study was to determine temporal dynamics of xylem and phloem formation in co-occurring deciduous and evergreen coniferous species in a dry inner Alpine environment (750 m a.s.l., Tyrol, Austria). By repeated micro-sampling of the stem, timing of key phenological dates of xylem and phloem formation was compared among mature Pinus sylvestris, Larix decidua and Picea abies during two consecutive years. Xylem formation in P. sylvestris started in mid and late April 2011 and 2012, respectively, and in both years about 2 week later in P. abies and L. decidua. Phloem formation preceded xylem formation on average by 3 week in P. sylvestris, and c. 5 week in P. abies and L. decidua. Based on modeled cell number increase, tracheid production peaked between early through late May 2011 and late May through mid-June 2012. Phloem formation culminated between late April and mid-May in 2011 and in late May 2012. Production of xylem and phloem cells continued for about 4 and 5-6 months, respectively. High variability in xylem increment among years and species is related to exogenous control by climatic factors and species-specific sensitivity to drought, respectively. On the other hand, production of phloem cells was quite homogenous and showed asymptotic decrease with respect to xylem cells indicating endogenous control. Results indicate that onset and culmination of xylem and phloem formation are controlled by early spring temperature, whereby strikingly advanced production of phloem compared to xylem cells suggests lower temperature requirement for initiation of the former.

Mismatch between primary and secondary growth and its consequences on wood formation in Qinghai spruce.

The connections between the primary and secondary growth of trees allows better understanding of the dynamics of carbon sequestration in forest ecosystems. The relationship between primary and secondary growth of trees could change due to the diverging responses of meristems to climate warming. In this study, the bud phenology and radial growth dynamics of Qinghai spruce (Picea crassifolia) in arid and semi-arid areas of China in 2019 and 2020 were weekly monitored to analyze their response to different weather conditions and their links with carbon sink. Xylem anatomical traits (i.e. lumen radial diameter and cell wall thickness) were quantified along cell radial files after the end of xylem lignification to calculate the early-to-latewood transition date. Winter and early spring (January-March) were warmer in 2020 with a colder April compared with 2019. Precipitation in April-June was lower in 2020 than in 2019. In 2019, bud phenology occurred earlier, while the onset of xylem formation and the early-to-latewood transition date were delayed. The duration from the beginning of split bud and exposed shoot to the early-to-latewood transition date was positively correlated with the radial width of earlywood (accounting for ~80% of xylem width) and total xylem width. The longer duration of xylem cell division did not increase xylem cell production and radial width. Moreover, the duration from bud burst to the early-to-latewood transition date in 2020 was negatively linked with early phloem cell production as compared with 2019. Our findings suggest that warm conditions in winter and early spring promote the xylogenesis of Qinghai spruce, but might delay bud burst. However, the xylem width increments largely depend on the duration from bud burst to the start of latewood cell division rather than on the earlier xylogenesis and longer duration of xylem cell differentiation induced by warm conditions.

Identification and Expression Profile of CLE41/44-PXY-WOX Genes in Adult Trees Pinus sylvestris L. Trunk Tissues during Cambial Activity.

WUSCHEL (WUS)-related homeobox (WOX) protein family members play important roles in the maintenance and proliferation of the stem cells in the cambium, the lateral meristem that forms all the wood structural elements. Most studies have examined the function of these genes in angiosperms, and very little was known about coniferous trees. Pine is one of the most critical forest-forming conifers globally, and in this research, we studied the distribution of WOX4, WOX13, and WOXG genes expression in Pinus sylvestris L. trunk tissues. Further, we considered the role of TDIF(CLE41/44)/TDR(PXY) signaling in regulating Scots pine cambial activity. The distribution of CLE41/44-PXY-WOXs gene expression in Scots pine trunk tissues was studied: (1) depending on the stage of ontogenesis (the first group of objects); and (2) depending on the stage of cambial growth (the second group of objects). The first group of objects is lingonberry pine forests of different ages (30-, 80-, and 180-year-old stands) in the middle taiga subzone. At the time of selection, all the trees of the studied groups were at the same seasonal stage of development: the formation of late phloem and early xylem was occurring in the trunk. The second group of objects is 40-year-old pine trees that were selected growing in the forest seed orchard. We took the trunk tissue samples on 27 May 2022, 21 June 2022, and 21 July 2022. We have indicated the spatial separation expressed of PsCLE41/44 and PsPXY in pine trunk tissues. PsCLE41/44 was differentially expressed in Fraction 1, including phloem cells and cambial zone. Maximum expression of the PsPXY gene occurred in Fraction 2, including differentiating xylem cells. The maximum expression of the PsCLE41/44 gene occurred on 27 May, when the number of cells in the cambial zone was the highest, and then it decreased to almost zero. The PsPXY gene transcript level increased from May to the end of July. We found that the highest transcript level of the PsWOX4 gene was during the period of active cell proliferation in the cambial zone, and also in the trees with the cambial age 63 years, which were characterized by the largest number of cell layers in the cambial zone. In this study, we have examined the expression profiles of genes belonging to the ancient clade (PsWOXG and PsWOX13) in stem tissues in Scots pine for the first time. We found that, in contrast to PsWOX4 (high expression that was observed during the period of active formation of early tracheids), the expression of genes of the ancient clade of the WOX genes was observed during the period of decreased cambial activity in the second half of the growing season. We found that PsWOX13 expression was shifted to Fraction 1 in most cases and increased from the phloem side, while PsWOXG expression was not clearly bound to a certain fraction. Based on the data, the role of the CLE41/44-PXY-WOX signaling module in regulating P. sylvestris cambial growth is discussed.