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.
Evaluating the state of deterioration of water-logged archaeological wood is necessary in order to select treatments for its conservation and storage, particularly in the case of valuable archaeological artefacts. For this purpose archaeological wood of ash (Fraxinus sp.) and oak (Quercus sp.) buried in water-logged conditions at prehistoric settlements on the Ljubljansko barje (Ljubljana moor), Slovenia, aged approx. 5,200 and 4,500 years, was investigated by means of light microscopy (LM), transmission electron microscopy (TEM) and cellular UV-microspectrophotometry (UMSP). LM and TEM revealed that the ash wood aged 5,200 years was the least preserved. The secondary walls of fibres, vessels and parenchyma cells were considerably thinner than in normal wood, indicating distinct degradation. TEM and UMSP additionally revealed strong delignification of the remaining parts of the secondary walls of all cell types. The compound middle lamellae appeared structurally intact, but had lower UV-absorbance than normal wood of the same species. The cell corners were topochemically unchanged, as shown by high analogue UV-absorbance. The UV-absorbance maxima at a wavelength of 278 nm corresponded to those of hardwood lignins. The oak heartwood was generally better preserved than the ash wood. Within each species, the 4,500- year-old samples generally appeared better preserved than those 5,200 years old.
Fir trees (Abies alba Mill.) in a permanently monitored forest in the Dinaric region in Slovenia respond to crown damages by distinctly reducing their ring widths. According to transmission electron microscopy (TEM) and UV-microspectrophotometry (UMSP) of cambium-adjacent latewood tracheids of affected trees, the secondary wall formation and lignification were completed by the middle of October. In samples taken at the same date from healthy looking silver firs, the S3 and the warty layer were not yet present in cambium-adjacent latewood tracheids. Additionally, their inner S2 showed lower lignin deposition, whereas the compound middle lamella, S1 and outer S2 were distinctly lignified as revealed by TEM and UMSP. It is assumed that these youngest tracheids of healthy trees will later undergo lignification or remain less lignified. From these observations we conclude that the cambial activity at breast height ends later in healthy silver fir trees as compared to declining trees.
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.
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.
The commonest approach to studying cambial productivity is conventional light microscopy, which is widely used in wood formation studies. The number of such studies has increased rapidly in the past decade, usually in order to elucidate the relationship between growth and environmental factors. However, some aspects of cambial seasonality are often overlooked or neglected. Observations with transmission electron microscopy provide a more detailed insight into changes occurring on the ultra-structural level in cambial cells. Criteria for defining cambial activity are not yet fully clarified, especially when observing it at different resolutions, i.e., on cellular, subcellular and ultrastructural levels. The goal of this review is to contribute to clarification of the terms mainly used, such as cambial dormancy, reactivation, activity, productivity and transition between different states, resting period and quiescence, which describe structural modifications of cambial cells during the various phases of their seasonal cycle. Based on our own cambium observations on adult beech trees growing at two different elevations, which were made with light and transmission electron microscopy, we discuss the influence of weather conditions on cambial activity and the advantage of the complementary use of different techniques and resolutions.
We studied the effect of local weather conditions on intra-annual wood formation dynamics and wood structure of European beech (Fagus sylvatica L.) from a temperate location in the Czech Republic in two consecutive years, 2010 and 2011, characterized by different amounts of precipitation. Microcores were taken at weekly intervals and transverse sections of cambial and xylem tissue were prepared for light microscopic observation. Air temperature and soil moisture content were measured daily at the research plot. Tree-ring formation patterns and vessel features showed different responses to climatic factors in the two years. In 2010, the onset of cambial cell production occurred almost 10 days later than in 2011, when a considerably reduced amount of rainfall was already observed in the winter and spring months, as shown in Standardized Precipitation Index (SPI) values. Lack of precipitation in 2011 caused premature cessation of cambial cell division and markedly narrower annual xylem increments. Vessel density and water conductive area were higher in 2011 than in 2010. Average vessel size in general did not change. In response to local weather conditions, beech controls its hydraulic conductivity mainly by changing the number of vessels and tree growth rate, followed by vessel size. The lower sensitivity of vessel diameter to hydrological alterations confirms previous studies by other authors.
Recent studies on the seasonal dynamics of secondary tissue formation in Mediterranean trees have shown that xylogenesis depends on species and site conditions, but many questions still remain open. On the other side of the cambium, even less information is available about phloem structure and timing of its formation. We analysed intra-annual phloem variation in width and cell traits in the conducting, non-collapsed phloem (CPH) of Pinus pinea and Pinus halepensis at Mediterranean sites in southern Italy and Spain. In all investigated trees, it was possible to differentiate among the non-conducting, collapsed phloem (NCPH), and the CPH. CPH showed no evident annual growth layers; no differences in radial dimensions of early- and late phloem sieve cells, and no cyclic patterns of axial parenchyma distribution. Since it was not possible to study the seasonality of the phloem growth, we analysed the entire CPH. CPH width showed seasonal fluctuations and was generally the widest during the maximum cambial activity and narrowest during summer and winter. The radial size of newly formed sieve cells varied in relation to seasonal dynamics of cambial activity and fluctuations in local weather conditions. The number of axial parenchyma cells in CPH increased during the summer. The observed intra-annual variations in CPH width and structure seemed to be correlated with seasonal weather conditions in order to ensure a sufficient amount of conducting phloem tissue for translocation of photosynthates and signalling molecules to the actively growing tissues along the stem of a tree growing in the harsh Mediterranean conditions.
The responses of the vascular cambium and tracheid differentiation to extreme drought in Aleppo pine (Pinus halepensis Mill.) were investigated. The research focused on the drought year of 2005, in the primary study area at Maigmo (MAI) in southeastern Spain, with comparisons in Jarafuel (JAL) and Guardamar (GUA). The climate in this region is typically warm and dry with hot summers. Wood formation throughout the 2005 growing season was studied in transverse microtome sections and integrated with a retrospective dendrochronological analysis of crossdated increment cores collected in 2009. For most anatomical sections collected throughout the growing season at MAI, the vascular cambium appeared to be dormant as indicated by the low number of cells per radial file. Occasionally, immature xylem derivatives were observed during the growing season but without production of an annual ring. In increment cores collected at MAI, the 2005 position in the annual ring series contained either a narrow ring of both earlywood and latewood (47% of samples), a narrow ring of apparent latewood with no earlywood (13%), or a missing ring (50%). We introduce the term “dark ring” to refer to those annual rings of apparent latewood with no earlywood. For trees at JAL, the 2005 ring had below-average width and contained both earlywood and latewood. At GUA, the trees produced the widest 2005 ring of all three sites and mainly contained an intra-annual density fluctuation (IADF). The IADF was formed after cambial reactivation in the autumn. Although dark rings, IADFs, and especially missing rings complicate dendrochronological analysis, these anatomical features may provide an additional proxy record from which to infer climate variability and change in the past.
Annual periodicity of cambium production of xylem and phloem cells has rarely been compared in trees from different environments. We compared the structure of cambium and the youngest xylem and phloem increments in four tree species, Fagus sylvatica, Picea abies, Pinus sylvestris and Pinus halepensis, from nine temperate and Mediterranean sites in Slovenia and Spain. In Picea abies, Pinus sylvestris and Fagus sylvatica from temperate locations in Slovenia, xylem and phloem growth ring boundaries could be identified. In Fagus sylvatica growing at two elevations on Moncayo mountain, Spain, phloem increment consisted of only early phloem. In Pinus sylvestris from the same two sites, growth ring boundaries were not as clear as in temperate Slovenian sites. In some cases we could identify phloem growth ring boundaries but in others it was very doubtful, which could be explained by collapse of the outermost early phloem sieve cells. In Pinus halepensis from all sites, we could only distinguish between collapsed and non-collapsed phloem, while phloem rings could not be identified. Widths of the youngest phloem and xylem annual increments could only be compared when phloem increments could be clearly defined, as with Picea abies, Fagus sylvatica and Pinus sylvestris from temperate sites. The visibility of the growth ring boundary in phloem was not related to the width of annual radial growth. The correlation between xylem and phloem ring widths was high, but moderate between the number of dormant cambial cells and xylem ring and phloem ring widths. Based on the structure of the youngest phloem increments, we concluded that there is no typical annual periodicity in cambial production of phloem cells in trees from certain Mediterranean sites. This may be due to continuous yearlong cell production and the absence of true cambium dormancy, at least on the phloem side, under mild winter conditions.