Ethylene evolution,radial growth and carbohydrate concentrations in Abies balsamea shoots ringed with Ethrel

The terminal (1-year-old) shoot of quiescent, 2-year-old Abies balsamea (L.) Mill. seedlings was either left untreated or ringed with 0, 1 or 10 mg Ethrel g(-1) lanolin. After 5 weeks of culture under environmental conditions favorable for growth, the shoots were harvested to measure ethylene evolution and carbohydrate concentrations by gas chromatography, and tracheid number and bark radial width by microscopy. In untreated shoots, the basal rate of ethylene evolution followed the order: cambial region > cortex + periderm = xylem + pith = needles. Wound-induced ethylene production was not detected until at least 4 h after excision, but was evident in all fractions 24 h after excision; the increase in wound-induced ethylene evolution followed the order: cambial region > cortex + periderm > xylem + pith > needles. Compared with untreated controls, the application of plain lanolin, which involved the removal of needles and periderm, increased bark radial width and wound-induced ethylene production by the cambial region and the cortex + periderm, but decreased cambial region concentrations of fructose, glucose and starch at the application point. At the application point, Ethrel concomitantly increased ethylene evolution from the cambial region and the cortex + periderm, tracheid number, bark radial width, and the cambial region concentrations of fructose, glucose, sucrose and starch. No effects of Ethrel treatment were detected above or below the application point, with the exception that the 10 mg g(-1) Ethrel treatment stimulated ethylene evolution and decreased starch concentration of the cambial region. The results indicate that: (1) the cambial region is the major source of endogenous ethylene in the 1-year-old shoot; (2) the magnitude of the difference in ethylene evolution between particular shoot fractions is different before and after the start of wound-induced ethylene production; (3) the Ethrel-induced increase in tracheid number and bark radial width at the application point is positively related to ethylene evolution from the cambial region and the cortex + periderm, respectively; and (4) ethylene derived from Ethrel applied laterally to a woody stem can mobilize carbohydrates to the application point.     

Laterally applied Ethrel causes local increases in radial growth and indole-3-acetic acid concentration in Abies balsamea shoots

The terminal (1-year-old) shoot of quiescent, two-year-old balsam fir (Abies balsamea (L.) Mill.) seedlings was ringed with lanolin containing 0, 1 or 10 mg g(-1) Ethrel, an ethylene-generating compound, and cultured for 6 weeks under environmental conditions favorable for growth. Bud break and the elongation of the current-year terminal shoot were monitored, and the subjacent previous-year terminal shoot that had been treated with Ethrel was harvested to measure stem radial growth by microscopy, shoot ethylene evolution by gas chromatography, and cambial region indole-3-acetic acid (IAA) concentration by combined gas chromatography-mass spectrometry. Compared with the lanolin controls, Ethrel at 1 and 10 mg g(-1) did not affect bud break or longitudinal growth, but stimulated tracheid production and bark increment up to about 2-fold at the application site, though not above or below it. In addition, the 1 and 10 mg g(-1) Ethrel treatments increased the cambial region IAA concentration about 3-fold and the evolution of ethylene at least 40-fold at the application site, compared with unwounded portions of both treated and control shoots. The 10 mg g(-1) Ethrel treatment also stimulated ethylene evolution about 10-fold, both above and below the application site. However, this stimulation was not associated with an elevation in cambial region IAA concentration. Similarly, the lanolin control treatment increased ethylene evolution at the application site about 10-fold, without affecting the cambial region IAA concentration. Our results suggest that the localized stimulation of radial growth in woody shoots ringed with Ethrel is mediated by an increase in IAA concentration, which in turn is induced by a threshold, abnormally high concentration of Ethrel-derived ethylene.     

Induction of acropetal (14)C-photosynthate transport and radial growth by indole-3-acetic acid in Pinus sylvestris shoots

The relationship between radial growth and assimilate movement was determined in one-year-old Pinus sylvestris (L.) cuttings collected during the dormant period and reactivated for 1-27 days under environmental conditions favorable for growth. The cuttings were either left with buds intact, defoliated along the distal 8 cm, or debudded and treated apically with 0 or 1 mg g(-1) indole-3-acetic acid (IAA) in lanolin. Radial growth was measured as tracheid production and bark radial width. The distribution of (14)C-photosynthate 1 or 5 days after exposure to (14)CO(2) was used to indicate assimilate movement. Debudding inhibited tracheid production and decreased acropetal (14)C-photosynthate transport, whereas applying IAA to debudded cuttings promoted both these processes and, in addition, induced vigorous callus growth within the bark immediately below the application point. Distal defoliation markedly increased the amount of (14)C-photosynthate transported toward the apex without altering how debudding and exogenous IAA affected tracheid production and the distribution of (14)C-photosynthate. The production of tracheids induced by apically applied IAA in debudded and distally defoliated cuttings increased progressively in material collected on September 9, October 14 and November 24, whereas bark radial width and acropetal (14)C-photosynthate movement increased only between the first two collection dates. Ringing midway between the IAA source and the bottom of the distally defoliated region with a lanolin mixture of 10 mg g(-1)N-1-naphthylphthalamic acid (NPA), 1 mg g(-1) methyl-2-chloro-9-hydroxy-fluorene-9-carboxylic acid (CF) or 10 mg g(-1) phenylacetic acid (PAA) reduced radial growth and [1-(14)C]IAA transport below the ring and locally promoted radial growth and accumulated radioactivity above. (14)C-Photosynthate transport into the region above the ring either was not altered (CF, PAA) or was increased (NPA). The results indicate that apically applied IAA induces the acropetal movement of (14)C-photosynthate in debudded P. sylvestris shoots by locally promoting activity correlated with tracheid and callus production rather than by affecting radial growth or phloem transport processes, or both, at a distance below the IAA source.     

Auxin and ethylene regulation of diameter growth in trees

Recent studies on the phytohormonal regulation of seasonal cell-division activity in the cambium, primary-wall radial expansion of cambial derivatives, differentiation of xylem cells, and growth of the cortex in forest trees of the north temperate zone are reviewed. Indol-3-ylacetic acid (IAA, auxin) has been characterized by combined gas chromatography-mass spectrometry (GC-MS) in the cambial region of Abies balsamea, Pinus densiflora, Pinus sylvestris and Quercus robur. All of the evidence supports the hypothesis that developing leaves and extending shoots are primary sources of IAA. The rate of ethylene emanation varies among conifer species when adjoining phloem and cambial tissues are incubated in vitro. The cambium from young cuttings of Abies balsamea produces more ethylene than that from older cuttings. Ethylene production by seven-year-old Abies balsamea cambium is substantially increased in vitro when the tissue is provided with exogenous 1-aminocyclopropane-1-carboxylic acid and IAA. In response to elevated ethylene concentrations, cortex growth is accelerated in both hardwood and conifer seedlings. Ethrel (2-chloroethylphosphonic acid) increases ray size and ray-cell number and promotes traumatic resin-canal development in xylem. In Ulmus americana, endogenous ethylene concentrations are inversely correlated with cambial activity. Ethylene decreases vessel diameter in Acer negundo, Acer platanoides and Ulmus americana. Several studies suggest that ethylene has a role in regulating reaction-wood formation in both conifers and hardwoods.     

Gravimorphism in current-year shoots of Abies balsamea: involvement of compensatory growth, indole-3-acetic acid transport and compression wood formation

We investigated the cause of gravimorphic growth inhibition in current-year shoots of balsam fir (Abies balsamea (L.) Mill.) seedlings displaced from their normal orientation in the gravitational field. Tilting the main stem of seedlings decreased shoot elongation, cambial growth as measured by tracheid production, and leaf dry weight of the terminal shoot and the lateral shoots on the lower side of the tilted stem. Removing either the terminal shoot or all lateral shoots induced compensatory growth in the remaining shoots, but did not reduce the inhibitory effect of tilting on shoot growth. Bending the apical part of a tilted main stem to restore it to the vertical did not fully reverse the inhibition of terminal shoot growth caused by stem tilting. Stem tilting inhibited cambial activity at the base of decapitated terminal shoots treated apically with indole-3-acetic acid (IAA) and decreased the basipetal transport of a [1-(14)C]-IAA pulse. Stem tilting also induced compression wood formation on the lower side of the tilted stem. Compression wood formation was associated with increases in cambial activity and stem respiration. Stem tilting did not affect either the net photosynthetic rate in 1-year-old leaves or the xylem water potential in current-year lateral shoots. These results support the hypothesis that gravimorphic growth inhibition in a current-year shoot on a tilted stem involves reductions in (1) the shoot's capacity to export IAA, and hence to mobilize photoassimilates, and (2) the supply of photoassimilates available for import by the shoot, as a result of increased cambial sink activity associated with compression wood formation outside that shoot.     

The relationship between crown size and ring width in Pinus sylvestris L. stems: dependence on indole-3-acetic acid, carbohydrates and nitrogen in the cambial region

Indole-3-acetic acid (IAA), carbohydrates, total nitrogen and amino acids in the cambial region and bark were measured at the top (10-year-old internode) and bottom (1.3 m) of the main stem of 50-year-old Pinus sylvestris L. trees, having different rates and longitudinal gradients of annual ring width formation. The trees were sampled during the most active period of wood production (June 23, July 15) and at the end of this period (August 23). Trees with a small crown and relatively slow growth rate (S-trees) were compared with trees in the same stand that had a large crown and fast growth rate (F-trees) as a result of thinning and fertilization. The effect of bottom pruning fast-growing trees (pruned F-trees) was also investigated. The F-trees had greater wood production than the S-trees at both the stem top and bottom. The difference was larger at the stem bottom, thus the relative decrease in ring width down the stem was steeper in the S-trees. The amount of sucrose and IAA per cm(2) in cambial region tissues was higher in F-trees than in S-trees. However, the differences in annual ring width between treatments and within trees were not reflected in the concentrations (expressed per gram fresh weight) of these substances, measured either in differentiating xylem, the cambium plus phloem, or in the cambial region as a whole. The concentrations of total nitrogen and amino acids were slightly higher in the F-trees than in the S-trees. Pruning the F-trees reduced wood production, particularly at the stem bottom, inhibited the springtime increase in starch, and decreased the amounts of sucrose and IAA per cm(2) in the cambial region. However, it was evident that the concentrations of sucrose, IAA, amino acid and nitrogen in the cambial region were not related to ring width. During June and July, the concentrations of these substances were generally higher at the stem bottom in the pruned F-trees than in the F-trees. The results provide evidence that it is the activity of the cambium rather than the availability of carbohydrates that determines the allocation of wood production along the tree stem. The results also indicate that tracheid production is not directly related to the IAA concentration in the cambial region.     

Control of longitudinal and cambial growth by gibberellins and indole-3-acetic acid in current-year shoots of Pinus sylvestris

We investigated the involvement of gibberellins (GAs) and indole-3-acetic acid (IAA) in the control of longitudinal and cambial growth in current-year shoots of Pinus sylvestris L. Elongating terminal shoots, located at the apex of previous-year (1-year-old) branches in the uppermost whorl on the main stem, were variously decapitated (apical 5 to 10 mm removed), defoliated (all developing needle fascicles removed) and treated with endogenous GA(4/7) or IAA, or both. Shoot length and the radial widths of xylem and phloem were measured, and the concentrations of GA(1), GA(3), GA(4), GA(9) and IAA in the stem were determined by combined gas chromatography-mass spectrometry with deuterated GAs and [(13)C(6)]-IAA as internal standards. Decapitation decreased the production of xylem and phloem and the IAA concentration, but did not alter either longitudinal growth or the concentrations of GAs. Defoliation markedly inhibited shoot elongation, as well as cambial growth, and reduced the concentrations of GA(1), GA(3), GA(4), GA(9) and IAA. Application of GA(4/7) to defoliated shoots promoted longitudinal growth and phloem production, without affecting xylem production or IAA concentration. Application of GA(4/7) and IAA together to decapitated + defoliated shoots increased shoot elongation, xylem and phloem production and IAA concentration, whereas applying either substance alone had a smaller effect or none at all. We conclude that, for elongating current-year shoots of Pinus sylvestris, (1) both the shoot apex and the developing needle fascicles are major sources of the IAA present in the stem, whereas stem GAs originate primarily in the needle fascicles, (2) GAs and IAA are required for both shoot elongation and cambial growth, and (3) GAs act directly in the control of shoot growth, rather than indirectly through affecting the IAA concentration.     

Seasonal variations of tracheid formation and amount of auxin (IAA) and gibberellin A4 (GA4) in cambial-region tissues of mature sugi (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis>) cultivar grown in a Nelder plot with different tree densities

The role of phytohormones in tracheid formation remains unclear in conifers. In this study, to obtain information on the role of auxin (IAA) and gibberellin A4 (GA4), we examined seasonal variation in the amount of phytohormones in cambial-region tissues and tracheid formation of sugi cultivar planted in a Nelder plot with different tree densities. We demonstrated that the amount of IAA was positively correlated with the number of tracheids formed in early and mid-season, but not in late season, and had no relation to tracheid differentiation. Crown length and height at the crown base had a positive and negative effect, respectively, on IAA amounts in early and mid-season, but not in late season. Height-to-diameter ratio was negatively correlated with IAA amounts in early and mid-season, but not in late season. Sugi trees with wider spacing continued tracheid formation in late season with smaller amounts of IAA, although the trees with narrower spacing ceased tracheid formation with larger amounts of IAA. Cambial growth cessation in late season might be controlled not by IAA amount, but by short-day-induced insensitivity to IAA. GA4 had no relation with the indexes of growth traits or tracheid formation.     

Variation in density of Picea sitchensis in relation to within-tree trends in tracheid diameter and wall thickness

The influences of cambial age and ring width on density of Sitkaspruce (Picea sitchensis (Bong.) Carr) were analysed in relationto within-tree trends in tracheid diameter and cell wall thickness.Discs were sampled at breast height from a total of 24 trees,from seven stands at three contrasting sites in Wales, and atbreast height, 30 per cent and 60 per cent total tree heightfrom one of the stands. Across the juvenile wood, ring density decreased with ring numberfrom the pith while radial tracheid diameter increased. Theseoverall trends were considered to be inherent to tree growth,presumably associated with cambial ageing, since they occurredin all trees on all sites. In juvenile wood, density also variedwith site growth rate (as indicated by ring width) at similarcambial age, wider rings being associated with more rapidrateof change in tracheid diameter with ring number and with decreasein tracheid wall thickness. Consequently, on a site having treeswith high growth rate density decreased more rapidly acrossthe juvenile wood, down to a lower minimum value, than on siteswith a slower growth rate. In mature wood, the decrease in densitywith increase in ring width was associated with differencesin both tracheid diameter and wall thickness. Density was slightly(though not significantly) higher at breast height than in comparablerings at 30 per cent total height, associated with significantlythicker tracheid walls at breast height. Changes in radial tracheid diameter (with ring number, or withring width) were associated with greater differences in theearlywood than towards the latewood end of each growth ring,while variations in wall thickness with ring width were associatedwith rate of increase in wall thickness towards the latewoodend. This may account for some previously conflicting reportson influence of silvicultural management on density, for densityis likely to vary with influence of environment on the seasonalcycle of cambial activity. The extent of the juvenile wood as delimited by the inner coreof wide growth rings does not necessarily correspond to theregion of varying tracheid dimensions in Sitka spruce.     

Identification and quantitation of indole-3-acetic and abscisic acids in the cambial region of Quercus robur by combined gas chromatography-mass spectrometry

A technique is described for the rapid purification of indole-3-acetic acid (IAA) and abscisic acid (ABA) from small quantities of the basal trunk cambial region tissues of 34-year-old trees of Quercus robur L. Mass spectra of IAA and ABA obtained from the purified extracts provide the first reported unequivocal identification of endogenous IAA and ABA in the cambial region of a hardwood species. By use of the combined gas chromatography-mass spectrometry technique of multiple ion monitoring, with deuterated analogs as internal standards, it was shown that the levels of IAA in the cambial region tissues varied from ~1 to ~20 microg g(-1) and those of ABA from ~1 to ~5 microg g(-1) dry weight during the growing season. The high levels of IAA are similar to those found in the newly differentiating xylem tissues of softwoods.     

乙烯与几种木本植物茎插穗生根的关系

生长素类物质(NAA,IBA,IAA)处理Ⅰ-69杨、香椿和枣的嫩枝插穗,可诱导乙烯释放;KT,ABA,GA_3处理则不能。生长素类物质处理插穗促进不定根形成,与诱导乙烯释放有关。在研究乙烯和乙烯利处理Ⅰ-69杨硬枝插穗对不定根形成作用中知得,低浓度促进,高浓度抑制。所以,不论是生长素类物质,还是乙烯或乙烯利,对不定根形成的促进或抑制作用,都与乙烯生成有关,是生长素与乙烯共同作用的结果.     

Tracheid length in relation to seedling height in conifers

Summary Changes in tracheid length during the development of primary and secondary xylem are analysed in relation to the division and elongation of cambial initials, in first year seedlings ofPinus sylvestris andPicea sitchensis. During primary and early secondary growth tracheid length varies with leaf internode length, but this relationship soon becomes obscured during later secondary growth. Correlations between seedling height and tracheid length thus depend on variation in tracheid length with radial xylem increment before and after terminal bud production. Since the relation between shoot and tracheid length is a complex one, the clonseness of the correlation varies with environmental treatment, but overall, a doubling of shoot length increased tracheid length by about 10%.I am grateful to ProfessorS. D. Richardson andDr. G. K. Elliott for many helpful suggestions. This work was financed by a grant from the Natural Environment Research Council. Plants were grown from seed kindly supplied by the Forestry Commission Research Division (Scotland and North England).     

植物生长调节剂对枸杞硬枝插穗发根及生长的影响

分别采用IAA、ABT生根粉,4种浓度水平,浸泡处理"宁杞1号"枸杞硬枝插穗进行扦插试验,结果表明:2种植物生长调节剂溶液质量浓度为100 mg.L-1,浸泡2 h对提高硬枝插穗成活率、促进枸杞扦插苗根、茎、叶的生长发育效果最好。     

毛白杨形成层的活动周期及其POD同工酶的变化

北京地区生长的毛白杨 (PopulustomentosaCarr .)在春季芽膨大前 ,形成层带已经进入了恢复活动期。在芽展开后一周形成层进入活动期 ,细胞开始分裂 ,细胞层数也有所增加 ,同时分化出未成熟的韧皮部细胞。未成熟韧皮部细胞的数量在形成层活动初期快速增长 ,随后是一个持续而缓慢的连续过程。木质部的产生比韧皮部要晚 3周 ,未成熟的木质部细胞层数在开始变化不大 ,5月底～ 6月底 ,增长速度明显高于初期 ,随后便急剧下降。成熟木质部细胞的出现比成熟的韧皮部细胞要晚约一个月。与其他研究结果不同 ,毛白杨在4月底到 6月底这段时间 ,形成层细胞层数迅速下降 ,未成熟韧皮部细胞也略有减少 ,而未成熟和成熟的木质部细胞则大量增加。木质部在 9月底停止分化 ,较韧皮部要晚约 2 0d。 9月底形成层进入休眠期后 ,形成层细胞层数基本保持不变 ,直到翌年春天恢复活动。毛白杨形成层区域的过氧化物酶 (POD)同工酶在形成层活动期其活性低、酶带少 ,而恢复活动期和休眠期其活性高且酶带多 ,组织化学反应还表明 ,不同时期POD在不同的细胞和组织中的活性不同。本文对POD同工酶在形成层活动周期中的变化与其组织分化的关系进行了讨论。     

Effects of mother tree ages,different rooting mediums,light conditions and auxin treatments on rooting behaviour of <Emphasis Type="Italic">Dalbergia sissoo</Emphasis> branch cuttings

Dalbergia sissoo Roxb. is one of the promising multipurpose tree species of South Asia. Most of the plantations of D. sissoo from seeds are facing severe threats due to the die-back disease, which ultimately causes death of this potential tree-species within a few months. Vegetative propagation could avoid the die-back disease. Thirty mother trees of different age-groups of D. sissoo were selected for evaluating the rooting behaviour of branch cuttings from D. sissoo as influenced by auxins (IAA or IBA at 100, 200, 500 mg·L⁻¹), ages of mother trees (10, 4 and 2 years old) and different environment conditions, i.e., different mediums (soil and sand) or light conditions (in shade and open condition). The results show that application of IAA and IBA induced more numbers of cuttings (collected from 10-year-old mother trees) to root compared to control. Branch cuttings of D. sissoo collected from 10-year-old mother trees and planted in soil bed in open conditions had 100.0% of cuttings to root in IAA (100 mg·L⁻¹) and IBA (200 mg·L⁻¹) treatments. Both rooting medium (Soil and sand) influenced significantly (p<0.05) on rooting response of branch cuttings. Soil medium was found to achieve maximum no. of branch cuttings to root, compared to sand medium.     

Effects of cutting types and hormonal concentration on vegetative propagation of Zanthoxylum armatum in Garhwal Himalaya,India

Experiments were carried out to define the effects of hormonal concentrations on semi-hard wood(SHW) and hard-wood(HW) branch cuttings of the Z.armatum. SHW and HW cuttings were collected in the month of March. The SHW and HW cuttings were treated with different concentration of indole-3-acetic acid and indole-3-butyric acid(IAA and IBA) and placed in vermiculite rooting medium for 90 days under 1-min misting after 10 min. Sprouting, rooting percentage, sprout number, sprout length, root number, and length were measured.The highest rooting and sprouting rate, 64.0 %, was obtained at the 0.3 % IBA treatment in the SHW cuttings.Similarly sprout length and number of roots per cutting were also higher at the 0.3 % IBA treatment in the SHW cuttings. The number of shoots per cutting was higher at the 0.3 % IAA treatment in the SHW cuttings. Root length per cutting was higher in 0.4 % IBA treatment in the SHW cuttings. The results indicated that 0.3 and 0.4 % IBA treatment produce higher rooting percentages as well as the number of roots and their length in SHW cuttings. The HW cuttings produced maximum rate of 18.0 % rooting in0.5 % IBA treatment. The control set and lower concentrations of IBA and IAA completely failed to root in the mist chamber.     

Identification of two compounds correlated with lack of rooting capacity of chestnut cuttings

Cuttings from mature chestnut (Castanea spp.) trees are very difficult to root, whereas juvenile or etiolated cuttings root easily. The HPLC chromatograms from mature cuttings consistently showed two well-defined peaks that were absent from juvenile or etiolated cuttings. The compounds responsible for these two peaks were isolated and identified as derivatives of ellagic acid, 3,3',4-tri-O-methylellagic acid (compound 1) and 3,4,4'-tri-O-methylellagic acid (compound II). Compound I significantly inhibited rooting as measured by the bean test; a concentration of 3.0 x 10(-5) M caused a 32% inhibition in number of roots formed compared with controls grown in 10(-5) M IAA. When compound I was mixed with an equal quantity of IAA, the rooting activity of the auxin was reduced by 31.5%. An insufficient amount of compound II was available for the bean assay.     

红毛五加扦插繁殖技术试验研究

通过对红毛五加穗条的不同药剂、扦插时间和扦插基质对红毛五加存活率影响的试验研究,结果表明：（1）红毛五加在河沙基质中扦插存活率最高,达78.75%;（2）ABT药剂50 mg.L-1、6 h处理穗条,2 a生枝条扦插存活率最高,达70%;（3）ABT、NAA、IAA、IBA药剂50 mg.L-1、6 h处理,比其他处理方式能显著提高扦插苗存活率。且ABT、NAA、IAA药剂50 mg.L-1、6 h处理可用于生产推广。     

几种柏科树种嫩枝扦插的研究

木本植物的插条繁殖至今仍是园林工作者获得良种无性系和培育苗木的重要途径。柏科树种多是珍贵的园林树种,它们的嫩枝插穗生根难易差别很大。本研究探讨了生根难易的生理原因,为提高扦插成活率提供理论依据。