Jožica Gričar

Seasonal Dynamics of Wood Formation in Pinus Halepensis from Dry and Semi-Arid Ecosystems in Spain

The seasonal dynamics 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 of RC in June or July. In general, tangential bands of RC 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.

Regular cambial activity and xylem and phloem formation in locally heated and cooled stem portions of Norway spruce

The effect of heating (23–25°C) and cooling (9–11°C) on regular cambial activity and xylem and phloem formation in the stem portion of Norway spruce was investigated. Adult trees were sampled at 21-day intervals during the 2005 vegetation period. Continuously elevated temperatures increased the rate of cell division in the first part of the growing season, but did not significantly prolong cambial activity at the end of the vegetation period in the heated tree. Low-temperature treatments shortened regular cambial activity and slowed down cell production. The xylem growth ring was wider in the heated sample and narrower in the cooled sample compared to the control. The temperature in the cambial region was only negligibly transferred along the stem from the site of its application. In general, the temperature in the cambium was affected by a long-term rise or drop in air temperatures. Both experiments affected the structure and width of phloem growth increments. The tangential band of the axial parenchyma was not continuous in the cooled sample. The number of late phloem cells was reduced in the cool-treated sample and increased in the heat-treated sample. Our experiments confirmed the effect of constantly increased or decreased temperatures on regular cambial activity in Norway spruce.

Woody biomass production lags stem-girth increase by over one month in coniferous forests

Wood is the main terrestrial biotic reservoir for long-term carbon sequestration(1), and its formation in trees consumes around 15% of anthropogenic carbon dioxide emissions each year(2). However, the seasonal dynamics of woody biomass production cannot be quantified from eddy covariance or satellite observations. As such, our understanding of this key carbon cycle component, and its sensitivity to climate, remains limited. Here, we present high-resolution cellular based measurements of wood formation dynamics in three coniferous forest sites in northeastern France, performed over a period of 3 years. We show that stem woody biomass production lags behind stem-girth increase by over 1 month. We also analyse more general phenological observations of xylem tissue formation in Northern Hemisphere forests and find similar time lags in boreal, temperate, subalpine and Mediterranean forests. These time lags question the extension of the equivalence between stem size increase and woody biomass production to intra-annual time scales(3, 4, 5, 6). They also suggest that these two growth processes exhibit differential sensitivities to local environmental conditions. Indeed, in the well-watered French sites the seasonal dynamics of stem-girth increase matched the photoperiod cycle, whereas those of woody biomass production closely followed the seasonal course of temperature. We suggest that forecasted changes in the annual cycle of climatic factors(7) may shift the phase timing of stem size increase and woody biomass production in the future.

Age dependence of xylogenesis and its climatic sensitivity in Smith fir on the south-eastern Tibetan Plateau.

An age effect on growth trends and climate/growth relationships of trees can possibly be discovered by analysing the seasonal dynamics of xylem development. The aims of this study, therefore, were to compare xylem formation of young (43 ± 4 years) and old (162 ± 26 years) Smith fir (Abies georgei var. smithii (Viguie & Gaussen) W. C. Cheng & L. K. Fu) trees in the Sygera Mountains, south-eastern Tibetan Plateau and, to identify the association between wood formation and climate. The seasonal radial growth dynamics of young and old trees was monitored on microcores collected at weekly intervals during two growing seasons. Transverse sections through phloem, cambium and outermost xylem of 9-12 μ m thickness were observed with a light microscope under bright field and polarized light to follow the cambial activity and differentiation of the developing xylem. Young trees were characterized by an earlier onset of xylogenesis, a longer growing season and a higher growth rate, resulting in a higher number of xylem cells. Both young and old trees responded fast to changes of the minimum air temperature, confirming that this factor was dominant by controlling Smith fir growth on the south-eastern Tibetan Plateau.

A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere.

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. METHODS 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. CONCLUSIONS 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.

The climate sensitivity of Norway spruce [Picea abies (L.) Karst.] in the southeastern European Alps

To investigate the potential of Norway spruce (Picea abies L. Karst) as a palaeoclimate archive in the southeastern European Alps, tree ring chronologies were developed from trees growing at two sites in Slovenia which differed in their ecological and climatological characteristics. Ring width, maximum latewood density, annual height increment and latewood cellulose carbon isotope composition were determined at both sites and the resulting time-series compared with and verified against instrumental climate data for their common period (AD 1960–AD 2002). Results indicate that ring width sensitivity to summer temperature is very site-dependent, with opposing responses at alpine and lowland sites. Maximum density responds to September temperatures, indicating lignification after cell division has ceased. Stable carbon isotopes have most potential, responding strongly to summer temperature in both alpine and lowland stands. Height increment appears relatively insensitive to climate, and is likely to be dominated by local stand dynamics.

Number of Cells in Xylem, Phloem and Dormant Cambium in Silver Fir (Abies Alba), in Trees of Different Vitality

We investigated the anatomical structure of phloem and xylem growth rings as well as the dormant cambium in relation to vitality in 81 adult silver fir trees (Abies alba Mill.). Specifically, we investigated the number of cells produced in the current phloem growth ring (PR), xylem growth ring (XR) and their ratio, the number of cells in the dormant cambium (CC), and the structure of the PR, which included characterisation of the early phloem (EP), the late phloem (LP), and the presence, absence, and continuity of tangential bands of axial parenchyma (AP). EP was relatively stable with respect to number and types of cells as PR width changed, but LP was quite variable. The CC of more vital trees produced more xylem than phloem cells. The ratio of XR to PR number decreased with decreasing vitality of trees and in the most severely affected trees (4% of the study group), more cells were formed in the PR than in the XR. The number of cells in phloem, xylem and dormant cambium is positively correlated. The use of width and structure of phloem and the ratios between PR, XR and CC can provide information on tree conditions and, consequently, can be a useful tool for forest management.

Plastic and locally adapted phenology in cambial seasonality and production of xylem and phloem cells in Picea abies from temperate environments

Despite its major economic importance and the vulnerability of Picea abies (L.) H. Karst. to climate change, how its radial growth at intra-annual resolution is influenced by weather conditions in forest stands with a high production capacity has scarcely been explored. Between 2009 and 2011, phenological variation in seasonal cambial cell production (CP) was analysed in adult P. abies trees from three contrasting sites, differing in altitude and latitude. The results indicate that the timing of cambial CP is a highly synchronic process within populations since in all cases the cambium simultaneously started and stopped producing xylem and phloem cells. Our results also demonstrate that the phenology of cambial CP is highly variable and plastic between years, depending on seasonal temperature and precipitation variation. Differences among sites, however, are only partially explained by different environmental (elevation and altitude) and climatic conditions, suggesting that local adaptation may also play a decisive role in the strategy of P. abies for adapting wood and phloem increments to function optimally under local conditions.

SeaSonal ultraStructural changeS in the cambial zone of beech (Fagus sylvatica) grown at two different altitudeS

Seasonal structural changes of cambial cells in mature beech (Fagus sylvatica L.) trees growing at elevations of 400 m a.s.l. (lowland) and 1200 m a.s.l. (mountains) are presented on the basis of light (LM) and electron microscopy (TEM). For LM, samples from trees were collected at weekly intervals and for TEM at two-month intervals from March till September, 2008. LM enabled us to follow the production of new xylem and phloem cells that lasted for 16.5 ± 3.7 weeks at the lowland site and for 10.7 ± 1.3 weeks in the mountains. TEM revealed differences in ultrastructure of cambial cells in the phases of dormancy, reactivation, activity and transition to dormancy. The seasonal patterns of ultrastructural changes in cambial cells were similar at both sites but their timing was different. TEM revealed changes in the fine structure of cambial cells, indicating their activation in spring and the earliest stages of cell divisions and development of new cell walls. When using LM, the onset of cambial activity could be observed one month later, compared with TEM. LM therefore enabled us to follow cambial productivity but not the activity and related cytoplasmic modifications during reactivation.

Topochemical investigations of cell walls in developing xylem of beech (Fagus sylvatica L.).

Abstract Formation and lignification of xylem cells in mature beech (Fagus sylvatica L.) trees growing in a forest site in Slovenia (46° N, 14°40′ E, 400 m a.s.l.) were studied on the cellular and subcellular level. Samples containing the cambial zone and developing xylem were taken from six beech trees every week throughout the 2006 vegetation period. Cell wall thickening and lignification in individual cell wall layers and cell types were determined by light microscopy, cellular UV-microspectrophotometry and transmission electron microscopy, respectively. Cell division started between the 18th and the 24th of April 2006. Lignification began in the newly formed xylem tissue on the 2nd of May. After 1 month, the developing earlywood portion contained fully differentiated vessels with completed wall deposition and lignification, and differentiated fibres and axial parenchyma became visible after 2 months. At the end of cambial cell division on the 9th August, the differentiation of the most recently formed fibres in the terminal zone of the growth ring continued for approximately 4 weeks. This indicates that the process of lignification in earlywood is slower than in latewood. The high temporal resolution of the investigated processes and the combination of the above-mentioned microscopic techniques provides a detailed insight into the process of cell wall thickening and lignification of woody tissue in beech.