Changes in the localization and levels of starch and lipids in cambium and phloem during cambial reactivation by artificial heating of main stems of Cryptomeria japonica trees

Background and Aims

Cambial reactivation in trees occurs from late winter to early spring when photosynthesis is minimal or almost non-existent. Reserve materials might be important for wood formation in trees. The localization and approximate levels of starch and lipids (as droplets) and number of starch granules in cambium and phloem were examined from cambial dormancy to the start of xylem differentiation in locally heated stems of Cryptomeria japonica trees in winter.

Methods

Electric heating tape was wrapped on one side of the stem of Cryptomeria japonica trees at breast height in winter. The localization and approximate levels of starch and lipids (as droplets) and number of starch granules were determined by image analysis of optical digital images obtained by confocal laser scanning microscopy.

Key Results

Localized heating induced earlier cambial reactivation and xylem differentiation in stems of Cryptomeria japonica, as compared with non-heated stems. There were clear changes in the respective localizations and levels of starch and lipids (as droplets) determined in terms of relative areas on images, from cambial dormancy to the start of xylem differentiation in heated stems. In heated stems, the levels and number of starch granules fell from cambial reactivation to the start of xylem differentiation. There was a significant decrease in the relative area occupied by lipid droplets in the cambium from cambial reactivation to the start of xylem differentiation in heated stems.

Conclusions

The results showed clearly that the levels and number of storage starch granules in cambium and phloem cells and levels of lipids (as droplets) in the cambium decreased from cambial reactivation to the start of xylem differentiation in heated stems during the winter. The observations suggest that starch and lipid droplets might be needed as sources of energy for the initiation of cambial cell division and the differentiation of xylem in Cryptomeria japonica.     

Relationship between the earlywood-to-latewood transition and changes in levels of stored starch around the cambium in locally heated stems of the evergreen conifer <Emphasis Type="Italic">Chamaecyparis pisifera</Emphasis>

Key message

We observed the formation of latewood tracheids with narrow diameters and thick walls and the disappearance of stored starch around the cambium on the locally heated region of stems in evergreen conifer Chamaecyparis pisifera during winter cambial dormancy.

Abstract

Wood formation is controlled by cambial cell division, which determines the quantity and quality of wood. We investigated the factors that control cambial activity and the formation of new tracheids in locally heated stems of the evergreen conifer Chamaecyparis pisifera. Electric heating tape was wrapped around one side of the stem, at breast height, of two trees in 2013 and two in 2014. Pairs of stems were locally heated in winter, and small blocks were collected from heated and non-heated regions of stems. Cambial activity and levels of stored starch around the cambium were investigated by microscopy. Cambial reactivation and xylem differentiation occurred earlier in heated than in non-heated regions. New cell plates were formed after 14–18 days of heating. After a few layers of tracheids with large diameters and thin walls had formed, cell division and cell enlargement during differentiation were inhibited. Tracheids with narrow diameters and thick walls, defining those as latewood, were formed near the cambium, and finally, four to six layers of tracheids were induced. After cambial reactivation, amounts of stored starch started to decrease and starch disappeared completely from phloem and xylem cells that were located near the cambium during the differentiation of heated regions. Our results suggest that an increase in temperature induces the conversion of stored starch to soluble sugars for continuous cambial cell division and earlywood formation. By contrast, a shortage of stored starch might be responsible for inhibition of cambial activity and induction of the formation of latewood tracheids.

     

A rapid decrease in temperature induces latewood formation in artificially reactivated cambium of conifer stems

Background and Aims Latewood formation in conifers occurs during the later part of the growing season, when the cell division activity of the cambium declines. Changes in temperature might be important for wood formation in trees. Therefore, the effects of a rapid decrease in temperature on cellular morphology of tracheids were investigated in localized heating-induced cambial reactivation in Cryptomeria japonica trees and in Abies firma seedlings. Methods Electric heating tape and heating ribbon were wrapped on the stems of C. japonica trees and A. firma seedlings. Heating was discontinued when 11 or 12 and eight or nine radial files of differentiating and differentiated tracheids had been produced in C. japonica and A. firma stems, respectively. Tracheid diameter, cell wall thickness, percentage of cell wall area and percentage of lumen area were determined by image analysis of transverse sections and scanning electron microscopy. Key Results Localized heating induced earlier cambial reactivation and xylem differentiation in stems of C. japonica and A. firma as compared with non-heated stems. One week after cessation of heating, there were no obvious changes in the dimensions of the differentiating tracheids in the samples from adult C. japonica. In contrast, tracheids with a smaller diameter were observed in A. firma seedlings after 1 week of cessation of heating. Two or three weeks after cessation of heating, tracheids with reduced diameters and thickened cell walls were found. The results showed that the rapid decrease in temperature produced slender tracheids with obvious thickening of cell walls that resembled latewood cells. Conclusions The results suggest that a localized decrease in temperature of stems induces changes in the diameter and cell wall thickness of differentiating tracheids, indicating that cambium and its derivatives can respond directly to changes in temperature.     

Differentiation of terminal latewood tracheids in silver fir trees during autumn

BACKGROUND AND AIMS: The differentiation of terminal latewood tracheids of silver fir (Abies alba) trees grown in Slovenia was investigated in autumn/winter 2001/2002. METHODS: The experimental trees were divided into three groups: one with narrow annual rings, width less than 1 mm; one with annual ring widths between 1 and 4 mm; and one group with broad rings larger than 4 mm. The differentiation of terminal latewood tracheids was investigated by light-, electron- and UV-microscopy in tissues sampled in October and November 2001 and March 2002. KEY RESULTS: In the middle of October, cambial divisions did not occur any more in any of the trees. In trees with narrow annual rings, cell wall deposition as well as lignification were completed in terminal latewood tracheids at this date, whereas in trees with annual ring widths of more than 1 mm these processes still continued. Electron microscopy as well as UV microscopy revealed an unlignified inner S(2) layer and the absence of S(3) and warty layers. With increasing distance from the cambium, wall formation and lignification gradually appeared to be completed. Samples of all trees taken in the middle of November only contained differentiated terminal latewood tracheids. At the structural and lignin topochemical level, November and March samples showed completed differentiation of walls of terminal latewood tracheids. CONCLUSIONS: In trees with broader annual rings, the final steps of differentiation of the youngest latewood tracheids near the cambium still continued during autumn, but were finished prior to winter. It was concluded from structural observations that duration of cambial activity is longer in trees with broad annual rings than in trees with narrow rings.     

Effect of local heating and cooling on cambial activity and cell differentiation in the stem of Norway spruce (Picea abies)

BACKGROUND AND AIMS The effect of heating and cooling on cambial activity and cell differentiation in part of the stem of Norway spruce (Picea abies) was investigated. METHODS: A heating experiment (23-25 degrees C) was carried out in spring, before normal reactivation of the cambium, and cooling (9-11 degrees C) at the height of cambial activity in summer. The cambium, xylem and phloem were investigated by means of light- and transmission electron microscopy and UV-microspectrophotometry in tissues sampled from living trees. KEY RESULTS: Localized heating for 10 d initiated cambial divisions on the phloem side and after 20 d also on the xylem side. In a control tree, regular cambial activity started after 30 d. In the heat-treated sample, up to 15 earlywood cells undergoing differentiation were found to be present. The response of the cambium to stem cooling was less pronounced, and no anatomical differences were detected between the control and cool-treated samples after 10 or 20 d. After 30 d, latewood started to form in the sample exposed to cooling. In addition, almost no radially expanding tracheids were observed and the cambium consisted of only five layers of cells. Low temperatures reduced cambial activity, as indicated by the decreased proportion of latewood. On the phloem side, no alterations were observed among cool-treated and non-treated samples. CONCLUSIONS: 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.     

Cambium Destruction in Conifers Caused by Pinewood Nematodes

Percentage and rate of mortality in 2-4-year-old conifers depended upon the numbers of pinewood nematodes Bursaphelenchus xylophilus inoculated into their stems. In addition, percentage of conifer mortality was greater for spring inoculations when cambial activity was greater than for late summer and fall inoculations. Gross and histological examination of stems revealed destruction of the cambial layer, including fusiform and ray intitials and their derivatives. These data suggest that cambial and ray destruction causes tree death through blockage of tracheids by gas, oleoresin, or metabolites from dying ray tissues.     

Concentrations of oxygen and indole-3-acetic acid in cambial region during latewood formation and dormancy development in Picea abies stems

To manipulate the occurrence of latewood formation and cambial dormancy in Picea abies (L.) Karst. stems, potted seedlings were transferred from the natural environment on 9 July, when tracheids early in the transition between earlywood and latewood were being produced, and cultured for up to 5 weeks in a controlled environment chamber having: (1) Warm LD, (25/15C during day/night) and long (16 h) photoperiod, (2) Warm SD, (25/15C) and short (8 h) photoperiod, or (3) Cold SD, (18/8°C) and short (8 h) photoperiod. In Warm LD trees, the radial enlargement of primary-walled derivatives on the xylem side of the cambium, as well as xylem production, continued at the same magnitude throughout the experiment. In Warm SD and Cold SD trees, the radial enlargement of primary-walled derivatives declined and the cambium entered dormancy, both developments occurring faster in the Warm SD trees. The concentrations of indole-3-acetic acid (IAA) was higher in developing xylem tissue than in cambium+phloem tissues, but did not vary with environmental treatment or decrease during the experimental period. The O2 concentration in the cambial region followed the order of Cold SD>Warm SD>Warm LD trees and was <5%, the threshold for the inhibition of IAA-induced proton secretion, for the first 3 weeks in Warm SD and Warm LD trees. Thus, neither latewood formation nor cambial dormancy can be attributed to decreased IAA in the cambial region. Nor does lower O2 concentration in the cambial region appear to be inhibiting the IAA action that is associated with cambial growth.     

Cavitation of intercellular spaces is critical to establishment of hydraulic properties of compression wood of Chamaecyparis obtusa seedlings

Background and Aims When the orientation of the stems of conifers departs from the vertical as a result of environmental influences, conifers form compression wood that results in restoration of verticality. It is well known that intercellular spaces are formed between tracheids in compression wood, but the function of these spaces remains to be clarified. In the present study, we evaluated the impact of these spaces in artificially induced compression wood in Chamaecyparis obtusa seedlings.Methods We monitored the presence or absence of liquid in the intercellular spaces of differentiating xylem by cryo-scanning electron microscopy. In addition, we analysed the relationship between intercellular spaces and the hydraulic properties of the compression wood.Key Results Initially, we detected small intercellular spaces with liquid in regions in which the profiles of tracheids were not rounded in transverse surfaces, indicating that the intercellular spaces had originally contained no gases. In the regions where tracheids had formed secondary walls, we found that some intercellular spaces had lost their liquid. Cavitation of intercellular spaces would affect hydraulic conductivity as a consequence of the induction of cavitation in neighbouring tracheids.Conclusions Our observations suggest that cavitation of intercellular spaces is the critical event that affects not only the functions of intercellular spaces but also the hydraulic properties of compression wood.     

Cambial sensitivity to rising temperatures by natural condition and artificial heating from late winter to early spring in the evergreen conifer Cryptomeria japonica

Differences in the timing of cambial reactivation and the initiation of xylem differentiation in response to the sum of daily maximum temperatures were studied in two Cryptomeria japonica trees with cambium of different ages under natural and locally heated conditions. In addition, we observed the effects of low temperature on cambial activity. The timing of cambial reactivation and of the initiation of xylem differentiation differed between 55- and 80-year-old cambium under natural conditions. In the 55-year-old cambium, cambial reactivation occurred when the cambial reactivation index (CRI), calculated on the basis of daily maximum temperatures in excess of 10°C, was 94 and 97°C in 2007 and 2008, respectively. In 80-year-old cambium, cambial reactivation occurred when the CRI, calculated on the basis of daily maximum temperatures in excess of 11°C, was 69 and 71°C in 2007 and 2008, respectively. After cambial reactivation in 2007, normal cell division was evident in the cambium even though the minimum temperature had fallen between −2 and −3°C. Under natural conditions, xylem differentiation started 38–44 days after cambial reactivation. In heated stems, the time between cambial reactivation and the initiation of xylem differentiation ranged from 14 to 16 days, a much shorter time than under natural conditions, indicating that continuous exposure to an elevated temperature had induced earlier xylem differentiation. Our observations indicate that the sensitivity to reactivation inducing stimuli of the cambium depends on both the stage of dormancy and tree age of the cambium.     

How needle phenology indicates the changes of xylem cell formation during drought stress in Pinus sylvestris L.

Temperature, water availability and photoperiod are the primary drivers of tree phenological processes. However, there is lack of information how the spring temperature and summer drought affect phenology of both cambium and needles. We evaluated the interplay between xylem and needle development of mature Scots pine trees in three consecutive years differing in the spring temperatures and onset and duration of summer drought. Cambial activity began on the day of a year (DOY) 83–87, while the bud break occurred on DOY 113–119, when also the secondary cell wall of tracheids started its formation. While the timing of bud break correlated with the sum of effective temperatures, the beginning of cambial zone activity did not. The needles were fully unfolded around DOY 170, which corresponded to the transition between earlywood and latewood tracheids. Summer drought did not affect needle development, but it changed the rate of production and morphology of latewood tracheids. Latewood tracheids from the year 2015 when the drought was longest (i.e. until the second half of August in 2015 compared to the duration from June until the middle of July in the two other years) were 32% narrower and they had 34% thinner cell walls than in 2014 and 2016. The improvement of tree water status in July resulted in the formation of intra-annual density fluctuations (IADF). The interplay between needle and cambium phenology provided an insight into how the tree allocates the resources with varying temperature and soil water availability.     

Induction of cambial reactivation by localized heating in a deciduous hardwood hybrid poplar (Populus sieboldii x P. grandidentata)

BACKGROUND AND AIMS: The timing of cambial reactivation plays an important role in the control of both the quantity and the quality of wood. The effect of localized heating on cambial reactivation in the main stem of a deciduous hardwood hybrid poplar (Populus sieboldii x P. grandidentata) was investigated. METHODS: Electric heating tape (20-22 degrees C) was wrapped at one side of the main stem of cloned hybrid poplar trees at breast height in winter. Small blocks were collected from both heated and non-heated control portions of the stem for sequential observations of cambial activity and for studies of the localization of storage starch around the cambium from dormancy to reactivation by light microscopy. KEY RESULTS: Cell division in phloem began earlier than cambial reactivation in locally heated portions of stems. Moreover, the cambial reactivation induced by localized heating occurred earlier than natural cambial reactivation. In heated stems, well-developed secondary xylem was produced that had almost the same structure as the natural xylem. When cambial reactivation was induced by heating, the buds of trees had not yet burst, indicating that there was no close temporal relationship between bud burst and cambial reactivation. In heated stems, the amount of storage starch decreased near the cambium upon reactivation of the cambium. After cambial reactivation, storage starch disappeared completely. Storage starch appeared again, near the cambium, during xylem differentiation in heated stems. CONCLUSIONS: The results suggest that, in deciduous diffuse-porous hardwood poplar growing in a temperate zone, the temperature in the stem is a limiting factor for reactivation of phloem and cambium. An increase in temperature might induce the conversion of storage starch to sucrose for the activation of cambial cell division and secondary xylem. Localized heating in poplar stems provides a useful experimental system for studies of cambial biology.     

Relationships between cambial activity,cell differentiation and the localization of starch in storage tissues around the cambium in locally heated stems of<Emphasis Type="Italic"> Abies sachalinensis</Emphasis> (Schmidt) Masters

A study was made, in a cool-temperate zone, of the extent of cell division in the cambium, the extent of differentiation of cambial derivatives, and the localization of storage starch around the cambium in locally heated (22–26°C) stems of the evergreen conifer Abies sachalinensis (Schmidt) Masters during cambial dormancy and immediately after natural reactivation of the cambium. In locally heated regions of stems during cambial dormancy, heating induced localized reactivation of the cambium. However, the cells in the heated and reactivated cambium stopped dividing soon after only a few cells had been generated. In addition, no differentiation of the xylem and the disappearance of starch from storage tissues around the cambium were observed. In regions of stem that had been locally heated after natural reactivation of the cambium, cell division continued in the cambium and earlywood tracheids with a large radial diameter and secondary walls were formed, with abundant starch in the storage tissues around the cambium. Our results suggest that the extent of both cell division in the cambium and cell differentiation depends on the amount of starch in storage tissues around the cambium in the locally heated stems of an evergreen conifer growing in a cool-temperate zone.     

Photosynthesis-nitrogen relationship in a Hinoki cypress (Chamaecyparis obtusa) canopy: a comparison with Japanese cedar (Cryptomeria japonica)

The relationship between light-saturated photosynthetic capacity (P _max) and leaf nitrogen (N) content was investigated for one year in a 15-year-old Chamaecyparis obtusa canopy and was compared with a Cryptomeria japonica canopy previously described. The linear regression between P _max and leaf N content tended to converge toward a single line segment from July to January and in May for C. obtusa. The slope of the linear regression between P _max and leaf N content of C. obtusa was gentler than that of C. japonica. The smaller regression coefficient of C. obtusa may reflect species differences in nitrogen nutrition requirements between C. obtusa and C. japonica. A pronounced decrease in the slope of the linear regression lines due to low temperature was observed in February and March. During this period, P _max of C. obtusa declined more than that of C. japonica suggesting that C. obtusa is less tolerant to low temperatures than C. japonica.     

Anatomical features that facilitate radial flow across growth rings and from xylem to cambium in Cryptomeria japonica

Background and Aims

Although the lateral movement of water and gas in tree stems is an important issue for understanding tree physiology, as well as for the development of wood preservation technologies, little is known about the vascular pathways for radial flow. The aim of the current study was to understand the occurrence and the structure of anatomical features of sugi (Cryptomeria japonica) wood including the tracheid networks, and area fractions of intertracheary pits, tangential walls of ray cells and radial intercellular spaces that may be related to the radial permeability (conductivity) of the xylem.

Methods

Wood structure was investigated by light microscopy and scanning electron microscopy of traditional wood anatomical preparations and by a new method of exposed tangential faces of growth-ring boundaries.

Key Results

Radial wall pitting and radial grain in earlywood and tangential wall pitting in latewood provide a direct connection between subsequent tangential layers of tracheids. Bordered pit pairs occur frequently between earlywood and latewood tracheids on both sides of a growth-ring boundary. In the tangential face of the xylem at the interface with the cambium, the area fraction of intertracheary pit membranes is similar to that of rays (2·8 % and 2·9 %, respectively). The intercellular spaces of rays are continuous across growth-ring boundaries. In the samples, the mean cross-sectional area of individual radial intercellular spaces was 1·2 µm² and their total volume was 0·06 % of that of the xylem and 2·07 % of the volume of rays.

Conclusions

A tracheid network can provide lateral apoplastic transport of substances in the secondary xylem of sugi. The intertracheid pits in growth-ring boundaries can be considered an important pathway, distinct from that of the rays, for transport of water across growth rings and from xylem to cambium.Key words: Cryptomeria japonica, bordered pit, intercellular spaces, lateral transport, tracheid network, water conduction, xylem permeability     

Amylostereum laevigatum associated with the Japanese horntail,Urocerus japonicus

The fungus associated with the Japanese horntail,Urocerus japonicus, in Kochi, Kagawa and Ehime Prefectures was studied. Cultures isolated from the mycangia of 113 adult females of the horntail showed the same cultural characteristics. Four of basidiocarps found on felled logs ofCryptomeria japonica were identifieds asAmylostereum laevigatum based on morphological characteristics. This was the first record ofA. laevigatum from Japan. The cultures isolated from the basidiocarps had the same cultural characteristics as those from the mycangia ofU. japonicus. One mycangial isolate produced basidiocarps on artificially inoculated stem segments ofCr. japonica after a 6-mo incubation and was identified asA. laevigatum. One isolate from the basidiocarps ofA. laevigatum and one from the mycangium ofU. japonicus were artificially inoculated into five trees each ofChamaecyparis obtusa andCr. japonica. The wood of all inoculated trees showed discoloration, with no difference in shape and pattern of discoloration between the two isolates. The inoculated fungi were reisolated from the areas of discoloration in the inoculated trees.     

Downhill movement of litter and its implication for ecological studies in three types of forest in Japan

The amount of litter moving down the slope was measured in three types of forest, together with an examination of rain as a factor in bringing this about. The three forest types were a natural mixed stand ofPinus densiflora and hardwood trees (plot A), aCryptomeria japonica plantation (plot S) and aChamaecyparis obtusa plantation (plot H). The amount of moved litter was quite large in plots A and H, but relatively small in plot S. The rain factor had little influence on litter movement in plot A, but was the main cause of movement in plot S and (especially) plot H. It is suggested that measurement of litter input and output not only by traps above ground level, but also by ones on the ground is essential for determining the cycling of elements inC. obtusa forests. It is also suggested that the decomposition of leaf litter should be studied both on the soil surface and in the soil inC. obtusa forests.     

Stem anatomy of Dolichos lablab Linn (Fabaceae): Origin of cambium and reverse orientation of vascular bundles

Secondary growth in the stem of Dolichos lablab is achieved by the formation of eccentric successive rings of vascular bundles. The stem is composed of parenchymatous ground tissue and xylem and phloem confined to portions of small cambial segments. However, development of new cambial segments can be observed from the obliterating ray parenchyma, the outermost phloem parenchyma and the secondary cortical parenchyma. Initially cambium develops as small segments, which latter become joined to form a complete cylinder of vascular cambium. Each cambial ring is functionally divided into two distinct regions. The one segment of cambium produces thick-walled lignified xylem derivatives in centripetal direction and phloem elements centrifugally. The other segment produces only thin-walled parenchyma on both xylem and phloem side. In mature stems, some of the axial parenchyma embedded deep inside the xylem acquires meristematic activity and leads to the formation of thick-walled xylem derivatives centrifugally and phloem elements centripetally. The secondary xylem comprises vessel elements, tracheids, fibres and axial parenchyma. Rays are uni-multiseriate in the region of cambium that produces xylem and phloem derivatives, while in some of the regions of cambium large multiseriate, compound, aggregate and polycentric rays can be noticed.     

Cambial reactivation in locally heated stems of the evergreen conifer Abies sachalinensis (Schmidt) Masters

A study was made of cambial activity, the localization of storage starch around the cambium, and the localization and occurrence of microtubules in cambial cells from dormancy to reactivation in locally heated (22–26 °C) stems of the evergreen conifer Abies sachalinensis. Heating induced localized reactivation of the cambium in the heated portions of the stem. Erect ray cambial cells resumed cell division 1 d prior to the reactivation of fusiform cambial cells and procumbent ray cambial cells. The re-initiation of the division of fusiform cambial cells occurred first on the phloem side. During the heat treatment, the amount of storage starch decreased in procumbent ray cambial cells and in the phloem parenchyma adjacent to the cambium but increased in fusiform cambial cells. Preprophase bands of microtubules, spindle microtubules and phragmoplast microtubules were observed both in erect ray cambial cells and in procumbent ray cambial cells. By contrast, no evidence of the presence of such preprophase bands of microtubules was detected in fusiform cambial cells. The results suggest that the localized heating of stems of evergreen conifers might provide a useful experimental model system for studies of the dynamics of cambial reactivation in intact trees. Received: 25 May 2000 / Accepted: 12 July 2000     

Cellular level observation of water loss and the refilling of tracheids in the xylem of Cryptomeria japonica during heartwood formation

The mechanism of heartwood formation in Cryptomeria japonica D. Don has long been studied since heartwood formation is a fundamental physiological feature of trees. In this study, the water distribution in the xylem of C. japonica was investigated at the cellular level to reveal the role of water distribution in the xylem during heartwood formation. Samples were taken from different heights of each trunk, in which the phases of heartwood formation differed. These were designated as SIH, which consisted of sapwood, intermediate wood, and heartwood; SI, which consisted of sapwood and intermediate wood but no heartwood; and S-all, which consisted entirely of sapwood. Cryo-scanning electron microscopic observations of the heartwood-formed (SIH) and non-heartwood-formed (SI and S-all) xylem revealed different patterns of water distribution changes in tracheids between the latewood and earlywood. In the latewood, almost all tracheids were filled with water in all areas from the sapwood to the heartwood (98–100% of tracheids had water in their lumina). In the earlywood, however, the water distribution differed between the sapwood (95–99%), intermediate wood (7–12%), and heartwood (4–100%). Many of the tracheids in the xylem, where the sapwood changed to intermediate wood lost water. In the heartwood, some tracheids remained empty, while others were refilled with water. These results suggest that the water distribution changes in individual tracheids are closely related to heartwood formation. Water loss from tracheids may be an important factor inducing heartwood formation in the xylem of C. japonica.     

Effects of environmental factors on wood formation in Scots pine stems

Summary To find the optimal conditions for growth and development of tracheid walls in Scots pine stems the effects of temperature and precipitation on xylem cell production by the cambium, radial cell expansion and secondary wall thickening have been studied. The observations were carried out on 10 specially chosen 50 to 60-year-old trees, growing in central Siberia, over 2 seasons. The data on the number of cells in differentiation zones and mature xylem along radial rows of tracheids, radial and tangential sizes of tracheids and their lumens were used for calculating cambial activity, the rates and durations of cell development in the zones, and both the thickness and cross sectional areas of tracheid walls. The mean day, mean maximal diurnal and mean minimal nocturnal temperatures have been shown by correlation and regression analyses to affect differentially separate stages of cytogenesis. The temperature influenced the initial division the side of xylem and radial cell expansion mainly in May–June, while the influence of precipitation increased in July–August. Throughout all seasons it was the temperature that had the main influence on the biomass accumulation in cell walls. Optimal values of temperature and precipitation for cell production by cambium, radial cell expansion and secondary wall thickening have been calculated. The data are discussed in connection with productivity and quality of wood.