Participation of auxin and abscisic acid in the regulation of seasonal variations in cambial activity and xylogenesis

Summary The current notion that hormonal level and cell response are clearly correlated has often been challenged recently. During the period of cambial activity, auxin content seems to control the intensity of mitosis and some features of the resulting wood, but not the duration of the active period itself. During cambial rest, the indole-3-acetic acid (IAA) level often remains high in the cambium, but the cell sensitivity to auxin is low. The decrease of auxin transport in autumn is sometimes interpreted as a major qualitative change affecting the pattern of transport, and sometimes as a secondary change occurring later than rest onset. The causes of the seasonal variation of cambial response remain unknown. A hypothesis is proposed that accounts for the structural-functional changes occurring in cambial cells during the onset of dormancy. Abscisic acid (ABA) may reduce wood production and xylem cell enlargement in late summer. An important amount of ABA may be present in the cambial zone in autumn after drought stress and in spring in the young growing shoot. Changes in ABA level do not appear to be clearly correlated with the different steps of cambial rest and activity. Beyond the role of ABA as a stress mediator, its participation in the annual regulation of cambial activity remains unclear. Its distribution in the most alkaline compartments may account for the particularities of its seasonal activity. The involvement of IAA and ABA in cambial growth is discussed within the scope of a possible annual alternation of two different metabolisms in the cambial cell.Abbreviations ABA abscisic acid - DPA dihydrophaseic acid - GA gibberellic acid - GC-MS gas chromatography-mass spectrometry - IAA indole-3-acetic acid - PA phaseic acid - RNA ribonucleic acid - SICM single ion current monitoring - SIM selected ion monitoring     

Level of endogenous indole-3-acetic acid in the stem of Pinus sylvestris in relation to the seasonal variation of cambial activity

Abstract. Gas chromatography – selected ion monitoring – mass spectrometry was used to measure the level of indole-3-acetic acid (IAA) in the cambial region at the top and bottom of the branchless portion of the main stem of three large Scots pine trees, at weekly intervals from 28 April to 13 July. During this period, the cambium reactivated from the dormant state and entered its 'grand' period of xylem and phloem production, which was monitored by microscopy. The total amount of IAA (ng cm⁻²) increased steadily from 28 April until late June, and thereafter remained constant. In contrast, the concentration of IAA (ng g⁻¹ fresh weight) was high at the start of cambial reactivation, declined when the number of differentiating tracheids began to increase, and then rose as the number of cells decreased. The timing and magnitude of the changes in xylem and phloem production and in IAA level were similar at the two sampling positions. It is concluded that the seasonal changes in cambial activity in the conifer stem cannot be ascribed simply to a fluctuation in the level of endogenous IAA in the cambial region.     

Inequality of size and size increment in Pinus banksiana in relation to stand dynamics and annual growth rate

BACKGROUND AND AIMS: Changes in size inequality in tree populations are often attributed to changes in the mode of competition over time. The mode of competition may also fluctuate annually in response to variation in growing conditions. Factors causing growth rate to vary can also influence competition processes, and thus influence how size hierarchies develop. METHODS: Detailed data obtained by tree-ring reconstruction were used to study annual changes in size and size increment inequality in several even-aged, fire-origin jack pine (Pinus banksiana) stands in the boreal shield and boreal plains ecozones in Saskatchewan and Manitoba, Canada, by using the Gini and Lorenz asymmetry coefficients. KEY RESULTS: The inequality of size was related to variables reflecting long-term stand dynamics (e.g. stand density, mean tree size and average competition, as quantified using a distance-weighted absolute size index). The inequality of size increment was greater and more variable than the inequality of size. Inequality of size increment was significantly related to annual growth rate at the stand level, and was higher when growth rate was low. Inequality of size increment was usually due primarily to large numbers of trees with low growth rates, except during years with low growth rate when it was often due to small numbers of trees with high growth rates. The amount of competition to which individual trees were subject was not strongly related to the inequality of size increment. CONCLUSIONS: Differences in growth rate among trees during years of poor growth may form the basis for development of size hierarchies on which asymmetric competition can act. A complete understanding of the dynamics of these forests requires further evaluation of the way in which factors that influence variation in annual growth rate also affect the mode of competition and the development of size hierarchies.     

Hormonal signals involved in the regulation of cambial activity, xylogenesis and vessel patterning in trees

The radial growth of plant stem is based on the development of cribro-vascular cambium tissues. It affects the transport efficiency of water, mineral nutrients and photoassimilates and, ultimately, also plant height. The rate of cambial cell divisions for the assembly of new xylem and phloem tissue primordia and the rate of differentiation of the primordia into mature tissues determine the amount of biomass produced and, in the case of woody species, the wood quality. These complex physiological processes proceed at a rate which depends on several factors, acting at various levels: growth regulators, resource availability and environmental factors. Several hormonal signals and, more recently, further regulatory molecules, have been shown to be involved in the induction and maintenance of cambium and the formation of secondary vascular tissues. The control of xylem cell patterning is of particular interest, because it determines the diameter of xylem vessels, which is central to the efficiency of water and nutrient transport from roots to leaves through the stem and may strongly influence the growth in height of the tree. Increasing scientific evidence have proved the role of other hormones in cambial cell activities and the study of the hormonal signals and their crosstalking in cambial cells may foster our understanding of the dynamics of xylogenesis and of the mechanism of vessel size control along the stem. In this article, the role of the hormonal signals involved in the control of cambium and xylem development in trees and their crosstalking are reviewed.     

Concomitant analysis of cambial abscisic acid and cambial growth activity in poplar

Endogenous levels of cambial region abscisic acid (ABA) were quantified by immunoassay and assessed together with cambial growth activity in poplar (Populus nigra L. × P. maximowiczii Henry, clone Kamabuchi) over the course of a growing season. The level of cambial region ABA increased from spring to late-summer but decreased sharply in autumn. Cambial growth activity, measured as the radial number of undifferentiated cambial cells and enlarging xylem cells, also increased from spring to summer and decreased sharply in autumn, indicating the onset of cambial dormancy. Exogenous ABA, applied laterally to poplar stems at two times within the growing season, enhanced cambial growth activity, as the radial number of undifferentiated cambial cells increased in ABA-treated trees subsequent to the two application times. Xylem cell development was also affected by exogenous ABA as fibre length increased significantly in ABA-treated trees at both application times. The positive correlation of cambial region ABA and cambial growth activity as well as the positive effects of exogenous ABA application thereon sheds new light on the role of this hormonal growth regulator.     

Patterns of inter-annual variation in the size asymmetry of growth in Pinus banksiana

A large body of literature suggests that asymmetric competition, where large individuals suppress the growth of smaller individuals by intercepting a disproportionate share of incoming light, is a dominant process in tree population development. This has not been examined extensively for long-lived tree species that accumulate growth over many years under varying growing conditions. Using dendrochronological techniques, we reconstructed annual growth and mortality rates at ten stands of jack pine (Pinus banksiana Lamb.) in Western Canada. We used these data to calculate an annual index of the size asymmetry of growth for each stand for the last 50 years. Jack pine is a shade-intolerant species found in even-aged monoculture stands, so the simple hypothesis is that large trees should consistently perform relatively better than small trees. Inter-annual variation in the index of size-asymmetric growth was positively associated with interannual variation in stand productivity at eight of ten sites. The size asymmetry of growth also showed a positive trend with age at eight of ten sites, even though all sites were in a period of declining leaf area. This should have reduced the intensity of asymmetric competition for light and reduced the size asymmetry of growth over time. Alternate hypotheses for this trend are (1) that physical collisions between crowns result in asymmetric competition for growing space because they are more damaging to small trees, or (2) that a differential build up of diseases in susceptible trees suppresses their growth, even in the absence of competition.     

Non-uniform distribution and seasonal variation of endogenous indol-3yl-acetic acid in the cambial region of Pinus contorta Dougl.

Endogenous, free indol-3yl-acetic acid (IAA) levels were measured in the main stem in the 10-year-old cambial zone, in the adjoining differentiating xylem, and in the adjoining mature xylem of 15–20-year-old Pinus contorta Dougl. by single-ion-current monitoring, combined gas chromatography — mass spectrometry, on several dates from early spring to early winter. Microscopy was used to determine the state of cambial activity on each harvest date. The IAA levels were found to be nearly constant at 1 g g^-1 DW in the cambial zone from March to July, then to increase to near 2 g g^-1 DW during the remainder of the growth season. No clear correlation was evident between number of fusiform cells per radial file and IAA content in the cambial zone. By contrast, the IAA content in differentiating xylem was higher than that in the adjoining meristematic zone on all harvest dates and also exhibited marked seasonal variation, peaking near 16 g g^-1 DW in mid summer, and declining to 1 g g^-1 DW in autumn. In mature xylem, IAA levels were very low and showed negligible variation. The fresh weight to dry weight ratio of differentiating xylem was greater than that of the cambial zone, and greater in the cambial zone than in mature xylem.     

Photosynthetic acclimation to elevated atmospheric carbon dioxide and UV irradiation in Pinus banksiana

Pinus banksiana seedlings were grown for 9 months in enclosures in greenhouses at CO₂ concentrations of 350 or 750 μmol mol⁻¹ with either low (0.005 to 0. 3 W m⁻²) or high (0.25 to 0. 90 W m⁻²) ultraviolet-B (UV-B) irradiances. Total seedling dry weight decreased with high UV treatment but was unaffected by CO₂ enrichment. High UV treatment also shifted biomass partitioning in favor of leaf production. Both CO₂ and UV treatments decreased the dark respiration rate and light compensation point. High UV light inhibited photosynthesis at 350 but not at 750 μmol mol⁻¹ CO₂ due to a UV induced increase in ribulose-1, 5-bisphosphate carboxylase/oxygenase efficiency and ribulose-1, 5-bisphosphate regeneration. Stomatal density was increased by high UV irradiance but was unchanged by CO₂ enrichment.     

Development of the Tapetum in Pinus banksiana Preceding Sporogenesis

Early in sporangial ontogeny, the cells destined to become thesporogenous and tapetal tissue differentiate in a strikinglysimilar manner. The first conspicuous step in development isa contraction of the protoplasts, beginning at the centre ofthe microsporangium and moving radially to its periphery. Similardevelopment of the two groups of cells ceases as the callosewall is formed around the meiocytes. At this point the originalwalls investing the tapetal cells become gelatinous, and lipidsynthesis commences within the contracted protoplasts. The bulkof this lipid is secreted from the cells, and becomes lodgedin the loculus, either as globules in the expanded radial andinner cell walls, or as a continuous layer on the inside ofthe middle lamella separating the loculus from the wall of themicrosporangium. This lipoidal layer forms the basement of aperitapetal membrane, believed to serve as a container for thefluid in which the young sporogenous cells are immersed. Examination of protein levels and ribosome numbers in the tapetalcells reveals that protein synthesis proceeds at an increasingrate throughout the development preceding meiosis, but apparentlyceases as the pollen mother cells become enveloped in callose.     

Proteolytic activity of clinical Candida albicans isolates in relation to genotype and strain source

Proteolytic activity is regarded as one of the most important virulence factors of Candida albicans. Several authors recently demonstrated that some karyotypes and genotypes harbouring a group I self-splicing intron (CaLSU) located in the gene encoding the large rRNA subunit showed a high level of proteinase production. The aim of this study was to investigate the correlation between the level of proteinase production and the presence of the CaLSU intron in C. albicans isolates originating from the blood and respiratory tracts (sputum/pharyngeal swabs) of patients with and without oropharyngeal candidosis. The results revealed statistically significant differences in genotype distribution and the level of proteinase production between the C. albicans isolates obtained from blood and from the respiratory tract. Genotype A, without the intron, was prevalent in all groups of strains and its prevalence was higher among isolates from blood (75%) and from patients with candidosis (80%) compared with strains from colonisation (as opposed to infection) (57.8%). Isolates from blood produced significantly less proteinase than isolates from the respiratory tract (p<0.02), and this difference should be attributed to lower proteinase production of genotypes B and C from blood compared with genotypes B and C from the respiratory tract (p<0.01). The higher proteinase production of genotype B than of genotype A was found among respiratory tract isolates only. The presented data indicate that the association between proteinase production and the CaLSU intron depends on the strains' population. Further study is needed on well-defined groups of clinical isolates to elucidate whether the observed diversity in proteinase production plays a role in the selection of strains inducing bloodstream infections.     

Malignant growth in relation to enzyme activity

Conclusion In this paper the recent work on the various phases of oxidation enzymes, glycolysis and metabolic radiation are coordinated.The cumulative evidence of this study as well as that of the specific functions of vitamins and hormones — all support the conception that malignant growth characterised by abnormal glycolysis and absence of mitogenetic radiation is primarily due to a protracted inhibition of the oxidation enzyme activity.This enzyme inhibition is brought about by (A) the depletion of the enzyme source and (B) environmental conditions suppressing normal enzyme activity.This dualism is however but apparent as in both cases the resulting abnormal metabolism leads to the synthesis of suboxidised glycolytic products, analogous, if not identical, to carcinogenic substances obtained by the pyrogenic dehydrogenation of sterols and sexual hormones.These active unsaturated compounds appear to disturb the sulphydryl equilibrium, connected with the glutathione function and essential to normal cell proliferation.The uniformity in the type of cancerous cell metabolism irrespective of the variety of external causes and applicable to both spontaneous and synthetic cancer — in other words regardless whether the carcinogenic bodies are introduced or producedin situ — supports this conception.     

Influence of exogenous ethylene on cambial activity,xylogenesis and ray initiation in young shoots of Leucaena leucocephala (lam.) de Wit

The effect of exogenous ethephon on cambial activity, xylem production and ray population in young shoots of Leucaena leucocephala was investigated anatomically. The application of ethephon showed a diphasic effect on cambial activity and xylogenesis in a dose dependent manner. Lower concentration of ethephon enhanced cambial activity while high concentrations reduced cambial cell divisions and daughter-cell differentiation. High ethephon concentration also resulted in shorter vessel elements, thick walled fibers and phenolic accumulation in ray cells and vessel elements, whereas low concentration allowed elongation of fibers and vessel elements. The density of rays increased significantly with increase in ethylene concentration. The evaluation of longitudinal sections of cambial zone in ethephon treated plants showed high frequency of transformation of fusiform initials into ray initials through divisions and segmentation, resulting in high ray frequency in both xylem and phloem. The present study demonstrates that ethylene plays an important role in regulating secondary vascular tissue composition by reducing the population of fusiform initials in the cambium.     

Cambial activity of Pinus elliottii var. densa reveals influence of seasonal insolation on growth dynamics in the Florida Keys

We determined the temporal and seasonal dynamics of intra-annual cell formation of south Florida slash pine (Pinus elliottii Engelm. var. densa Little & Dor.), the southernmost native pine in the United States and the foundation species of globally endangered pine rockland ecosystems. To assess intra-annual cambial activity and identify possible relationships between cell production and climatic factors, wood micro-cores were extracted monthly from six trees during the period March 2010 to March 2011. The results confirmed annual growth ring formation in P. elliottii var. densa and indicated that its growing season extends from February to December, with a short period of dormancy that varied little between individuals. Within the growing season, earlywood cells were produced from February to July, latewood cells produced from July to December, and intra-annual density fluctuations (IADFs) occurred in the growth rings of four of six trees between July and August. A principal component analysis indicated a homogeneous response of cambial activity among trees to site-specific climatic factors. The first principal component axis explained 71?% of the total variance in cell production during the study period. Our results indicated that the dynamics of seasonal cambial activity of P. elliottii var. densa are controlled by solar radiation (r?=?0.51, p?<?0.10) in the Florida Keys. The nature of our data allow us to only speculate on the ecophysiological processes responsible for IADFs in P. elliottii var. densa, and additional research is needed to better understand the relationship between their formation and the environment in the Lower Florida Keys.     

The seasonal variation in silicon concentration in chalk-streams in relation to diatom growth

SUMMARY. Changes in silicon concentration were used to investigate the seasonal cycle of diatoms in chalk-streams. Bore-hole water shows no significant variation in silicon concentration. Five kilometres downstream there is a significant reduction between March and May. Further downstream the cumulative effect of biological silicon removal shows more complex patterns. There is a considerable variation in the seasonal pattern from year to year but an analysis of the seasonal pattern averaged over thirteen years on the River Frome confirms a later summer silicon reduction in August-September, in addition to a much larger reduction in the spring.
The negative correlation between suspended chlorophyll-α and silicon concentrations confirms that suspended chlorophyll-α concentrations respond more closely to changes in the loosely attached diatom flora than to other sources of chlorophyll-α. However, neither variable is always suitable for monitoring changes in benthic diatoms. In headwaters, for example, transitory changes in diatoms do not have a significant impact on silicon concentration, while further downstream the suspended solids are affected by many other factors.
Estimates of diatom production, based on changes in silicon concentration, gave rates of 1–2 g Si m_-2 day_-1 in a small headwater chalkstream during the spring.     

Periodicity of cambial activity in Abies balsamea. I. Effects of temperature and photoperiod on cambial dormancy and frost hardiness

The relationship between from hardiness and growth potential, and their dependence on temperature and photoperiod, was investigated in the one-year-old cambium of balsam fir [Abies balsamea (L.) Mill.]. Six-year-old trees were exposed for 9 weeks to either the natural environment or one of 4 controlled environments in the fall (18 September-18 November), spring (12 April–14 June) and summer (19 July – 19 September). The 4 controlled environments were (1) WS, warm temperature (24/20°C in day/night) + short day (8 h). (2) WL. warm temperature (24/20°C) + long day (8 h + 1 h night break), (3) CS. cold temperature (9/5°C) + short day (8 h) and (4) CL, cold temperature (9/5°C) + long day (8 h + 1 h night break). At the beginning and end of each exposure, cambial activity was measured by recording the number of xylem, cambium and phloem cells, frost hardiness was estimated from the cambium's ability to survive freezing to –40°C, and cambial growth potential was deduced from the duration of the cell cycle and the production of xylem, cambium and phloem cells in cuttings cultured for 4 weeks with exogenous indole-3-acetic acid (IAA) under environmental conditions favourable for cambial activity. In the natural environment, frost hardening began in September and was completed in November, while dehardening occurred when the cambium reactivated. CL, CS, and to a lesser extent WS, promoted hardening in the summer and fall, but did not prevent dehardening in the spring. The cambial growth potential in the natural environment declined from a maximum in April to a low level in June, reached a minimum in September, then increased to a high level in November. This potential was promoted by CL and CS on all dates by WL in the summer and fall. The ratio of xylem to phloem induced by IAA treatment was greatest in June and least in September in cuttings from trees exposed to the natural environment, and was increased by CL and CS in the fall. The cambium in intact branches of trees protected from chilling during the fall and winter resumed cell cycling after less than 9 weeks of dormancy, but produced mostly or only phloem in the subsequent growing period. It is concluded that the frost hardiness of the cambium, the IAA-induced cycling of cambial cells, and IAA-induced xylem to phloem ratio vary independently with season, temperature and photoperiod, and that the periodicity of these processes is regulated endogenously.     

The original basal stem section influences rooting in Pinus banksiana cuttings

Photosystem II (PSII) from Cu-deficient pea plants ( Pisum sativum L., cv. Lincoln) has been investigated for electron transport activity, Cu content, and changes in some lipid components. Total fatty acid content was lower that in control plants, with an additional shift in the C₁₈ fatty acid patterns. Less α-linolenic and more linoleic and oleic acids were found. PSII preparations from Cu-depleted plants showed a decreased carotenoid content in light harvesting chlorophyll a/b complex of photosystem II (LHCII) and additional variations in pigment composition of pigment-protein complexes. In the green alga Dunaliella the effect of Cu deficiency on fatty acid composition was similar to that in pea plants, but the influence on the carotenoid pattern was much less pronounced.     

Proteolytic activity of vitreous-humour.

It has been found that the bovine vitreous--humour did not digest in vitro collagen at physiological and acidic pH. A proteolytic activity against haemoglobin in acid pH was found both in bovine and human vitreous-humours. The activity of human vitreous-humour increased significantly in endophtalmitis and glaucoma. In all pathological conditions studied the pH optima were at more acidic values than in control.