Localized cooling of stems induces latewood formation and cambial dormancy during seasons of active cambium in conifers

Background and Aims In temperate regions, trees undergo annual cycles of cambial growth, with periods of cambial activity and dormancy. Environmental factors might regulate the cambial growth, as well as the development of cambial derivatives. We investigated the effects of low temperature by localized cooling on cambial activity and latewood formation in two conifers, Chamaecyparis obtusa and Cryptomeria japonica.Methods A plastic rubber tube that contained cooled water was wrapped around a 30-cm-wide portion of the main stem of Chamaecyparis obtusa and Cryptomeria japonica trees during seasons of active cambium. Small blocks were collected from both cooled and non-cooled control portions of the stems for sequential observations of cambial activity and for anatomical measurements of cell morphology by light microscopy and image analysis.Key Results The effect of localized cooling was first observed on differentiating tracheids. Tracheids narrow in diameter and with significantly decreased cambial activity were evident 5 weeks after the start of cooling in these stems. Eight weeks after the start of cooling, tracheids with clearly diminished diameters and thickened cell walls were observed in these stems. Thus, localized low temperature induced narrow diameters and obvious thickening of secondary cell walls of tracheids, which were identified as latewood tracheids. Two months after the cessation of cooling, a false annual ring was observed and cambium became active again and produced new tracheids. In Cryptomeria japonica, cambial activity ceased earlier in locally cooled portions of stems than in non-cooled stems, indicating that the cambium had entered dormancy sooner in the cooled stems.Conclusions Artificial cooling of stems induced latewood formation and cessation of cambial activity, indicating that cambium and its derivatives can respond directly to changes in temperature. A decrease in the temperature of the stem is a critical factor in the control of cambial activity and xylem differentiation in trees.     

Callus Formation and Differentiation at an Exposed Cambial Surface

The origin of callus from the exposed surface of the cambialzone in Trema orientalis Bl. and Julbernardia globiflora (Benth.)Troupin is described and discussed in relation to what has beenobserved in other species. It is suggested that in differentspecies callus may develop from any of the undifferentiatedcentripetal products of the vascular cambium, but that the kindof tissue contributing to callus initiation depends upon thespecies and on the histology of the cambial zone. The establishment of a new vascular cambium and phellogen isalso described. It is found that cambium formation is not dependenton the presence of a preexisting cambium in the surroundingtissues and that the first-formed xylem is abnormal in structure.     

Seasonal Variation in the Ultrastructure of the Cambium in New Zealand Grown Pinus radiata D. Don

Examination of Pinus radiata cambium from trees growing in thecentral North Island of New Zealand has revealed that many ofthe structural changes occurring in other tree species at theonset or cessation of cambial activity are not found in thisspecies. The winter cambium bears a closer resemblance to thesummer cambium than it does to the winter cambium of any otherangiosperm or gymnosperm described in the literature. In particularthere is little change in vacuolar structure, endoplasmic reticulumform, and dictyosomal activity during the year. The only changeswhich take place involve a slight increase in vacuole volume,storage of starch in vacuoles, and a decrease in numbers ofspherosomes during the summer. These observations confirm that,while the degree of cambial activity is reduced in the winter,complete dormancy is absent.     

Cambial activity and annual rhythm of xylem production of Pinus kesiya Royle ex. Gordon (Pinaceae) in relation to phenology and climatic factors growing in sub-tropical wet forest of North East India

The interrelationship between phenological events, climatic factors, periodicity of cambial activity and seasonal production of xylem was examined in Pinus kesiya Royle ex. Gordon growing in sub-tropical wet forest of Meghalaya state, India. Reactivation of dormant cambium occurs after sprouting of new needles during the middle of February. Since the formation of reproductive cones takes place simultaneously with vegetative bud break and needle formation, cone formation could also lead to the enhancement of cambial activity. The activity of cambium and xylem production decline gradually towards November and cease from end of December to end of January. There was no correlation between needle fall and cambial activity. Due to the production of three flushes of new needles and branches in a year the tree never becomes completely leafless. It was evident from correlation and regression analysis that the annual course of average temperature plays an important role for the reactivation of vascular cambium after dormancy. The differentiation of xylem elements correlated with mean temperature in the first place and secondly with precipitation. Increase in length of fusiform initials and their derivatives could be correlated with relative humidity, precipitation and mean maximum temperature. Dormancy was imposed by low temperature and less precipitation. The data are discussed in the light of cambial activity, xylem production and phenological events.     

Seasonal Production of Secondary Phloem in the Twigs of Certain Tropical Timber Trees

Secondary phloem production in four deciduous (Albizzia lebbeck,Dalbergia sissoo, Tectona grandis and Terminalia crenulata)and three evergreen plants (Calophyllum inophyllum, Mangiferaindica and Morinda tinctoria) is briefly described. The totalduration of phloem production for each year was worked out forall these plants. In three of the four deciduous trees therewere two instalments of phloem production in correspondencewith the presence of two flushes of cambial activity while inTectona grandis and in all the three evergreen trees there wasonly one instalment. The time of initiation and cessation ofphloem tissue production was found to be variable in the differentplants studied. Periodicity in the production of different componentsof phloem tissue as well as the difference in the dimensionsof the different phloic elements produced during each flushof cambial activity resulted in detectable growth increments(or ‘rings’) within the phloem. There was a distinctshortening of the different phloem elements during the approachof dormancy/least activity. Conspicuous changes were found inthe ergastic contents of phloem parenchyma and ray cells adjacentto the cambial zone during the initiation of cambial activityand during the approach of dormancy/least activity. Seasonal growth, secondary phloem, deciduous and evergreen trees, cambial activity     

Epicormic Bud Development in Quercus robur L. Studies of Endogenous IAA, ABA, IAA Polar Transport and Water Potential in Cambial Tissues10

Seasonal measurements of IAA,³ made using GC-MS, ⁴ indicatedthat in Q. robur the spring initiation of cambial activity andonset of visible bud outgrowth in the canopy is preceded byan increase in cambial region IAA. The effects of notch-girdlescut into the bole indicated that IAA in the cambial region laterwas present in separate physiological pools, with only the polar-transportedfraction affecting epicormic bud outgrowth. The stage in thespring when the epicormic buds grew out coincided with an increaseboth in cambial region IAA and in the capacity of cambial explantsfor IAA polar transport. Thus the stimulus needed by the epicormicbuds to overcome inhibition by polar-transported IAA appearedto be self-generated. The observed effects of exogenous hormoneson epicormic bud outgrowth from stem explants indicated thatthis stimulus might be cytokinin. The seasonal changes detectedin cambial region ABA³ were consistent with a role for stress-inducedABA in the induction of epicormic bud dormancy after canopydevelopment during the summer. No consistent effects of standthinning on cambial region IAA, ABA, water potentials or watercontents were detected, although polar transport of exogenousIAA by cambial region explants removed in the spring was reducedby thinning. Key words: Epicormic buds, cambium, hormones     

Cambial activity and annual rhythm of xylem production of elephant apple tree (<Emphasis Type="Italic">Dillenia indica</Emphasis> Linn.) in relation to phenology and climatic factor growing in sub-tropical wet forest of northeast India

The interrelationship between phenological events, climatic factors, periodicity of cambial activity and seasonal production of xylem was examined in Dillenia indica L. (Dilleniaceae) growing in sub-tropical wet forest of Meghalaya state, India. The reactivation of cambial activity was seen in the first week of May, 15 days after sprouting of new leaves and buds. The activity of cambium and xylem production gradually declined toward December and ceased from January to April end. There was correlation between leaf fall and cambial dormancy. It was evident from the correlation and regression analysis, the relationship between cambial activity, xylem production with climatic factors, the monthly mean minimum temperature plays an important role for the cambial activity and xylem production rather than influence by rainfall and relative humidity in D. indica L. The data were discussed in the light of cambial activity, xylem production and phenological events.     

Cambial Activity in Acacia raddiana Savi

Cambial activity and extension growth of Acacia raddiana Saviwere investigated. Both types of growth are strongly dependenton relatively high temperatures. In contrast with the behaviourof the cambium of trees with a clear interrupted growth period,the cambium of A. raddiana saplings did not respond to the twodifferent photoperiods tried. A correlation between extension growth and cambial activitycould not always be observed. The cambium was always activeonly in young green twigs, and in their close proximity, underconditions where intensive extension growth took place. In oldertwigs the cambial activity was found to occur independentlyof the extension growth, even in the very same branches in whichextension growth occurs.     

Cambial Activity in Zygophyllum dumosum Boiss

Cambial activity in Zygophyllum dumosum Boiss.—a dominantdesert shrub in Israel—was investigated. Cambial activityin young plants seems to be greatly affected by water supply,while cambial activity in old specimens was found to be moreendogenously controlled. A late-summer cambial dormancy wasobserved under field conditions even in plants that were irrigated.     

Structural Changes in the Vascular Cambium of Pinus strobus L. during an Annual Cycle

The changes in the ultrastructure of cambial cells of easternwhite pine (Pinus strobus L.) during an annual cycle are observedand recorded as are relationships of cambial cells during dormancyand at resumption of cambial activity. Cambial activity wasresumed late in March or early in April, when a few cells dividedpericlinally. Cambial activity reached a maximum during thelatter part of May with 15 to 20 undifferentiated cells present.In July it declined markedly, and the number of undifferentiatedcells equalled that of the dormant period. The xylem and phloemtissue cells produced late in the annual cycle overwinteredat varying developmental stages. In October cambial cells structurallyresembled dormant cells. The number of dormant cells in easternwhite pine cambium varied from 6 to 10. Active cells were characterizedby a large central vacuole, by an abundance of all cell organelles,and by thin cell walls. Dormant cells were characterized bynumerous small vacuoles, by structurally and quantitativelymodified cell organelles, and by relatively thick cell walls.     

Seasonal Rhythms of Structure and Behaviour of Vascular Cambium in Ficus rumphii

Cambial structure and activity of Ficus rumphii Blume vary withthe changes in local climate. The cambial cells start swellingearly in April prior to the onset of periclinal divisions whichare most frequent in August. Cell division stops in October.During the growth season, initiation as well as cessation ofthe phloem production precedes that of xylem. A moderately hightemperature is correlated with the cambial reactivation. Onceinitiated, the activity continues at relatively low temperatures.Hot and dry environment favours the phloem production, whereashot and moderately humid conditions induce xylogenesis. Thesize and relative proportion of cambial initials also changewith season. Fusiform initials are shorter and broader duringthe rainy season (July–September) than for the rest ofthe year. Multiseriate and triseriate rays, as also the tallrays, outnumber the other types of rays throughout the year. Ficus rumphii, vascular cambium, phenology, climatic variation     

Variations in the Lengths of Fusiform Cambial Cells and Vessel Elements in Kalopanax pictus

Samples of a mature specimen of Kalopanax pictus, a ring-poroushardwood, were studied to compare the respective lengths offusiform cambial cells and vessel elements in the stem. Thelengths of dormant and reactivated fusiform cambial cells weremeasured with a confocal laser scanning microscope in tissuethat had been macerated by digestion with pectinase and in thicktangential sections. The lengths of early wood and late woodvessel elements were measured in tissues that had been maceratedby Franklin's method. The vessel elements and fusiform cambialcells varied considerably in length within individual samples.The mean length of early wood vessel elements corresponded tothat of fusiform cells in the dormant cambium but not in thereactivated cambium. Significant differences were observed betweenthe mean lengths of dormant and reactivated fusiform cambialcells, between those of reactivated fusiform cambial cells andearly wood vessel elements, between those of reactivated fusiformcambial cells and late wood vessel elements, and between thoseof early wood and late wood vessel elements. The frequency distributionsof lengths of cambial cells were bimodal and differed from thoseof vessel elements, which more closely resembled a normal distribution.The proportion of shorter lengths was higher in the reactivatedcambium than in the dormant cambium, the early wood and thelate wood vessel elements. Our results do not suppot the hypothesisthat the lengths of early wood vessel elements in ring-poroushardwoods change during differentiation. The similar rangesof recorded lengths suggest that short and long vessel elementsmight be derived directly from short and long cambial cells,respectively. Copyright 1999 Annals of Botany Company Kalopanax pictus, cambium, vessel element, confocal laser scanning microscopy, maceration, cell length.     

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     

毛白杨维管形成层带细胞超微结构的季节性变化

木材(次生木质部)是树木形成层细胞分化的产物,形成层的活动方式不仅影响木材的产量,而且影响木材的结构和性质.利用透射电子显微镜观察了生长在北京地区的毛白杨(Populus tomentosa Carr.)枝条形成层带细胞一个完整活动周期的超微结构变化.在木质部母细胞完全恢复活动之前,形成层纺锤状原始细胞的分裂和韧皮部细胞的分化已经开始.枝条上芽的展开和幼叶的生长可能决定了形成层带细胞的这种活动方式.透射电镜观察更清楚地揭示了树木形成层细胞在活动初期的分化特点.活动期形成层细胞中的大液泡在进入休眠期后逐渐分成许多小液泡分散在细胞质中.随着液泡融合逐渐消失的深色蛋白类物质又重新充满了大部分液泡.油滴和淀粉颗粒的年变化情况同液泡中的蛋白类物质基本相似.无论在活动期还是休眠期,形成层纺锤形细胞的质膜上都发现有许多可能与物质运输有关的小泡状内折.由核膜、内质网和高尔基体及其分泌小泡组成的细胞内膜系统,在形成层活动周期的不同阶段,其形态和分布明显不同,尤其在形成层细胞的恢复活动及其衍生木质部细胞次生壁的沉积过程中发挥着重要作用.整个活动周期中,形成层纺锤形细胞的径向壁都比弦向壁厚,处在休眠期的形成层带细胞,其径向壁与弦向壁的差别则更明显.形成层恢复活动时,径向壁上特别是与弦向壁相连的角隅处出现部分自溶现象.细胞壁特别是径向壁的变薄是形成层细胞恢复活动的重要特征.     

Opposite patterns in the annual distribution and time-course of endogenous abscisic acid and indole-3-acetic acid in relation to the periodicity of cambial activity in Eucommia ulmoides Oliv

The seasonal change of free abscisic acid (ABA) and indole-3-acetic acid (IAA) and their relationship with the cambial activity in Eucommia ulmoides trees were investigated by ABA and IAA immunolocalization using primary polyclonal and rhodamine-red fluorescing secondary antibodies, ABA and IAA quantification using high performance liquid chromatography (HPLC), and systematic monitoring of vascular cell layers production. ABA and IAA clearly displayed opposite annual distribution patterns. In the active period (AP), both immunolocalization and HPLC detected an abrupt decrease of ABA, reaching its lowest level in the summer. During dormancy, ABA started increasing in the first quiescence (Q1) (autumn), peaked in the rest (winter), and gradually decreased from the onset of the second quiescence (Q2) (the end of winter). IAA showed a reverse pattern to that of ABA: it sharply increased in AP, but noticeably decreased from the commencement of Q1. Longitudinally, the ABA distribution increased apico-basally, contrasting with IAA. Laterally, most of the ABA was located in mature vascular tissues, whereas the IAA essentially occurred in the cambial region. The concomitant IAA-ABA distribution and seasonal changes in vascular tissues greatly correlated with xylem and phloem cell production, and late wood differentiation and maturation. Interestingly, the application of exogenous ABA to quiescent E. ulmoides branches, in a water-culture system, inhibited external IAA action on cambial activity reactivation. These results suggest that, in E. ulmoides, ABA and IAA might probably interact in the cambial region. The annual cambial activity could be influenced by an IAA:ABA ratio; and ABA might play a key role in vascular cambium dormancy in higher plants. The relationship between hormonal changes and the (particular) annual life cycle of E. ulmoides is also discussed.     

Proteolytic activity in relation to seasonal cambial growth and xylogenesis in Pinus banksiana.

Proteolytic activity in the cambial zone and developing xylem of Pinus banksiana Lamb. was investigated over an annual cycle of growth and dormancy. Highest proteolytic activity was associated with the most active period of primary-wall radial expansion of cambial derivatives, in early spring, before protoplasmic autolysis was initiated in developing earlywood. Three pH maxima of proteolytic activity, near pH 3.0, 6.5 and 9.5, were observed at that time. In general, activities measured at pH values below 7.0 were greater than those determined above pH 7.0 at all stages in the annual cycle, in both cambial zone and developing xylem, although elevated activity at alkaline pH was also observed during springtime growth. Polyvinylpolypyrollidone (PVP) treatment markedly enhanced pH 7.5 but not pH 4.0 proteolytic activity in the cambial zone, but not in developing xylem, indicating the presence of PVP-binding proteinase regulators in the cambium. By fractionation and effector studies total proteolysis was determined to comprise interactions between serine, cystine, aspartate and metallo-proteases having MWs, by gel chromatography, between 10 and 100 kDa. The observations point to a complex regulatory mechanism controlling the presence and catalytic rates of the distinct types of proteases in the cambial region throughout an annual cycle of growth and dormancy.     

Distribution of Indole-3-Acetic Acid and the Occurrence of Its Alkali-Labile Conjugates in the Extraxylary Region of Pinus sylvestris Stems

Free and conjugated indole-3-acetic acid (IAA) were measured by quantitative gas chromatography-selected ion monitoringmass spectrometry in the extraxylary region of the stem of large Pinus sylvestris (L.) trees during the annual cycle of cambial activity and dormancy. The extraxylary region at the stem top and bottom was divided into 3 and 4 fractions, respectively, for the free IAA measurements, while the entire extraxylary region was extracted when the IAA-conjugates were analyzed. The effect on the distribution pattern of expressing IAA level as a concentration (per gram fresh weight or dry weight) and as total amount (per square centimeter) was examined. The IAA level was much higher in the cambial region than in the fractions that contained the nonfunctional phloem and the periderm. The largest IAA concentration occurred in the fraction that included the cambium, whereas the total amount of IAA was greatest in the phloemcontaining fraction. The significance of the nonuniform radial distribution of IAA for estimating the IAA concentration in the cambial region is discussed in relation to how the cambial region is sampled. A slight Iongitudinal gradient in IAA concentration, decreasing from the top to the bottom of the stem, was observed in the cambial region when the cambium was in the grand period of activity, but not at the end of the cambial growing period. In all fractions, the total amount of IAA was highest when the cambium was active. However, the IAA concentration in the cambial region did not follow the same pattern, actually being lowest during the tracheid production period at the stem bottom. IAA conjugates were detected on all sampling dates except June 23, but their concentrations were always less than 14% of that of free IAA, and their occurrence did not obviously vary during the year. In general, there was a higher concentration of ester conjugates than of amide conjugates, and the ester conjugates were more abundant at the top of the stem than at the bottom.     

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.     

Seasonal Changes in the Effects of Auxin on Rooting in Stem Cuttings of Ficus infectoria

Ficus infectoria stem cuttings were treated with 10 and 100 μg/ml each of IAA, IBA, 2,4, -D and NAA at monthly intervals and planted to study their rooting response after recording morphophysiological status and cambial activty of the parent branches. Attempts were also made to surgically expose the cambium before auxin treatment to determine the relationship of seasonal variation in auxin effectivity to cambial activity. The results show that: (1) there are two distinct phases in the sensitivity of Ficus infectoria stem cuttings to auxin-induced rooting; (2) the high rooting phase coincides with renovation of growth and high cambial activity starting in March and lasting through August and the low rooting phase coincides with winter dormancy and low cambial activity; (3) roots emerge in longitudinal rows in slitted auxin-treated cuttings; (4) slitted auxin-treated cuttings root profusely in June when cambial activity is high but not in October when cambial activity is low suggesting a close correspondence of seasonal variation between the rooting activity of auxin and cambial activity.     

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.