Interaction between indole-3-acetic acid and ethylene in the control of tracheid production in detached shoots of Abies balsamea

We investigated the interaction between indole-3-acetic acid (IAA) and ethylene in the regulation of the seasonal periodicity of tracheid production in 1-year-old balsam fir (Abies balsamea (L.) Mill.) cuttings collected at different times during the dormant period. The cuttings were left with their buds intact or were debudded and treated either apically with IAA or 2-chloroethylphosphonic acid (Ethrel) in lanolin, laterally with IAA or Ethrel in lanolin, or basally with Ethrel, Co(2+) or Ag(+) in deionized water. The treated cuttings were then cultured for up to 5 weeks under controlled environment conditions favorable for cambial growth. No change in ethylene evolution was detected during the rest-quiescence transition, when IAA-induced tracheid production increased. The induction of cambial reactivation by IAA was associated with a rise in ethylene evolution, but there was no consistent relationship among IAA concentration, tracheid number and ethylene emission. Neither Ethrel, Co(2+) nor Ag(+) affected tracheid production when applied basally, except for 10 and 100 micro M Ethrel and 100 micro M Co(2+), which were inhibitory. In contrast, ethylene evolution was promoted by Ethrel and inhibited by Co(2+), whereas Ag(+) had no effect. Similarly, applying Ethrel apically or laterally increased ethylene evolution, but did not promote tracheid production except in the treatment in which 1 mg Ethrel g(-1) lanolin was applied laterally to cuttings treated apically with 0.1 mg IAA g(-1) lanolin, and in the treatment in which 10 mg Ethrel g(-1) lanolin was applied laterally to budded cuttings. We conclude that (1) ethylene evolution is not specifically associated with IAA-induced tracheid production, (2) ethylene does not mimic the promoting effect of IAA on tracheid production, and (3) ethylene can promote tracheid production, but only when its application results in a localized unphysiologically high concentration in the cambial region, which, in turn, induces an accumulation of IAA.     

Ethylene production during symptom development of pine-wilt disease

Ethylene release from Pinus thunbergii Parl. seedlings was examined after inoculation with virulent and avirulent isolates of Bursaphelenchus xylophilus. Enhanced ethylene release was observed only in seedlings inoculated with the virulent isolate. The avirulent isolate induced necrosis of xylem ray cells and embolism in some tracheids, however, the cambium and outermost xylem was intact where water conduction was maintained. The ethylene increase coincided with cambial death in seedlings inoculated with the virulent isolate.     

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.     

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.     

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.     

Influence of ectomycorrhizal fungal inoculation on growth and root IAA concentrations of transplanted conifers

We determined whether in vitro plant growth regulator production by mycorrhizal fungi is correlated with conifer seedling growth and root IAA concentrations. Container-grown seedlings of interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), lodgepole pine (Pinus contorta Dougl.) and ponderosa pine (Pinus ponderosa Dougl.) were inoculated at seeding with ectomycorrhizal fungi having a high, moderate or low capacity to produce either IAA or ethylene in vitro. Inoculated seedlings were grown for one season in the nursery, harvested in December, cold stored over winter and then transplanted to either a nursery field or a forest site in the spring. Seedling morphology and endogenous IAA in roots were measured immediately after cold storage and again six and 12 months after transplanting. Morphological responses to inoculation varied among different mycorrhizal fungi. Free IAA concentration of roots was increased in some inoculation treatments for all conifer species. In seedlings transplanted to a nursery field, in vitro ethylene-producing capacity of the ectomycorrhizal fungi was highly correlated with more morphological features than in vitro IAA-producing capacity. Both IAA- and ethylene-producing capacity were significantly correlated with more morphological features in seedlings transplanted to a forest site than in seedings transplanted to a nursery field. One year after transplanting, only in vitro IAA-producing capacity was correlated with endogenous IAA concentration of roots of the inoculated seedlings. We conclude that growth responses of conifer seedlings can be partially influenced by IAA and ethylene produced by ectomycorrhizal fungal symbionts.     

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。     

章古台造林树种引种与乡土树种对比栽培试验

为了选择造林树种,营造针阔混交林,在辽宁省章古台试验地,开展了造林树种引种与乡土树种对比栽培试验。试验表明:针叶树以樟子松、彰武松,其次是油松、东宁赤松的适应性比较强,生长量比较高;阔叶树以白榆、怀槐、刺榆、暴马丁香、桑树和色木槭的抗性比较强,生长比较快。     

Locating the initial in the vascular cambium of Pinus strobus L. by electron microscopy

Summary The results of this investigation by electron microscopy on the vascular cambium in Pinus strobus L. confirm the results of earlier investigations by light microscopy. First, they showed that differences in the thickness of tangential walls of cambial cells exist and that these differences permit grouping of cells according to the sequence of the previous divisions of the initial. This, in turn, permits the site of the initial cell in the cambial zone to be deducted. The thicker distal tangential wall of the initial in the successive series of tissue production allows for precluding the direction in which the initial function proceeds, toward the xylem or toward the phloem. Second, it was demonstrated that immature xylem cells exist in groups of four, that immature phloem cells exist in pairs, and that at all times more cells are produced toward the xylem than toward the phloem. Third, it was shown that the extra-thick tangential wall in immature xylem serves as a landmark to signal the changeover in the initial function from phloem to xylem production.     

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.     

秤锤树插穗生根的解剖学观察

从解剖学角度研究了秤锤树插穗生根的机制.结果表明:插穗扦插前在木质部、韧皮部、形成层以及皮层等部位都不存在潜伏根原基,根据生根类型的划分,秤锤树不定根应属于诱生根原始体.不定根根原基可由形成层细胞、愈伤组织细胞分化而来,逐渐突破皮层,伸向外部.     

Canopy gap characteristics and its influence on the regeneration of broad-leaved Korean pine forests in Changbai Mountain

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Physiological process of the symptom development and resistance mechanism in pine wilt disease

Pathophysiological changes during the symptom development of pine wilt disease are reconsidered from recent investigations. The symptom development is divided into two stages: the early and the advanced stages. In the early stage, small number of nematodes migrate in cortex, then in xylem of the stem, and induce denaturation and necrosis of parenchyma cells. These changes in parenchyma are regarded as defense reactions of pines which result in terpene synthesis in xylem cells and embolism in tracheids. Such changes in the early stage can be induced in both susceptible and resistant pine species by either virulent or avirulent isolates of pinewood nematode (Bursaphelenchus xylophilus), or byB. mucronatus. No change occur in physiological status of leaves, and nematode reproduction is suppressed during this stage. Pine trees can survive if symptom does not progress from this stage. The symptoms of the advanced stage usually occur only in susceptible pines infected by virulent nematode isolates. At the beginning of the advanced stage, enhanced ethylene production by stem which coincides with cambial destruction occurs, and results in embolism of the outermost xylem in the portion. The embolism causes decrease in leaf water potential and cessation of photosynthesis. After cessation of photosynthesis, symptoms develop drastically with a burst of nematode population. There seems to be some unknown mechanism which suppress nematode reproduction and invasion to the cambial zone. This mechanism is thought to be photosynthesis-dependent, so that in photosynthesis-decrased conditions, even avirulent nematodes can multiply and invade cambium to induce tree death. Water stress in hot and dry summer should accelerates symptom development from the early to the advanced stage through such decrease of photosynthesis-dependent “cambial resistance”.     

Hormonal physiology of wood growth in willow (Salix viminalis L.): effects of spermine and abscisic acid

Summary Willow cuttings from two-year-old twigs were grown in nutrient solution alone, nutrient solution with 0.1 mM abscisic acid or with the spermine. Roots and leaves emerged within three weeks and the cambium was activated in the shoot. In most cases earlywood was generated, even when the seedlings were made in late summer. In contrast to plantlets grown in regular nutrient solution, those treated with hormones either inhibited (ABA) or advanced (spermine) the formation of roots, leaves and wood. In addition, SEM observations of wood were combined with autoradiographic studies and metabolite analysis.¹⁴C-labeled photoassimilates from the leaves were unloaded from the phloem of the shoots and transported via the rays into the cambial zone and the xylem. In spermine treated plants labeled assimilates were highly concentrated in all cells of the newly built xylem. However, cells from plants treated with abscisic acid appeared only weakly labeled. Quantitative analysis of the assimilates after two-dimensional thin-layer chromatography showed that wood from spermine-treated plants accumulated 19% more assimilates than the control, while xylem from ABA-treated plants imported 81% less labeled compounds from the phloem. Thus, the results strongly support the view that hormones play a key role in wood formation.     

Changes in levels of endogenous plant hormones in cambial regions of stems ofLarix kaempferi at the onset of cambial activity in springtime

The total amounts of endogenous indole-3-acetic acid (IAA), cytokinins, and abscisic acid (ABA) were quantified by gas chromatography-selected ion monitoring-mass spectrometry (GC-SIM-MS) in cambial regions of the main stems ofLarix kaempferi during the spring season. During the sampling period, cambium in the dormant state entered the active meristematic state. The total amount of IAA did not change at the onset of cambial reactivation but increased when the active division of cambial cells became apparent. Four cytokinins —trans- andcis-ribosylzeatin (RZ),N ⁶-isopentenyladenine (iP),N ⁶-isopentenyladenosine (iPA) — were quantified, but no zeatin (Z) was detected. The total amount of the four cytokinins together and the total amount of isopentenyl-type cytokinins (iP and iPA) varied during the sampling period but did not appear to be specifically associated with cambial activity. The total amounts oftrans- andcis-RZ remained relatively constant during the sampling period, as did the total amount of ABA. The results suggest that there is little correlation between total amounts of endogenous plant hormones in the cambial region and reactivation of the cambium during the spring.     

Life strategies in intra-annual dynamics of wood formation: example of three conifer species in a temperate forest in north-east France

We investigated whether timing and rate of growth are related to the life strategies and fitness of three conifer species. Intra-annual dynamics of wood formation, shoot elongation and needle phenology were monitored over 3 years in five Norway spruces (Picea abies (L.) Karst.), five Scots pines (Pinus sylvestris L.) and five silver firs (Abies alba Mill.) grown intermixed. For the three species, the growing season (delimited by cambial activity onset and cessation) lasted about 4 months, while the whole process of wood formation lasted 5-6 months. Needle unfolding and shoot elongation followed the onset of cambial activity and lasted only one-third of the season. Pines exhibited an 'extensive strategy' of cambial activity, with long durations but low growth rates, while firs and spruces adopted an 'intensive strategy' with shorter durations but higher growth rates. We estimated that about 75% of the annual radial increment variability was attributable to the rate of cell production, and only 25% to its duration. Cambial activity rates culminated at the same time for the three species, whereas shoot elongation reached its maximal rate earlier in pines. Results show that species-specific life strategies are recognizable through functional traits of intra-annual growth dynamics. The opposition between Scots pine extensive strategy and silver fir and Norway spruce intensive strategy supports the theory that pioneer species are greater resource expenders and develop riskier life strategies to capture resources, while shade-tolerant species utilize resources more efficiently and develop safer life strategies. Despite different strategies, synchronicity of the maximal rates of cambial activity suggests a strong functional convergence between co-existing conifer species, resulting in head-on competition for resources.     

Cambial Anatomy and Annual Rhythm of Secondary Xylem Development in the Twigs of Azadirachta indica A. Juss. (Meliaceae) Growing in Different Forests of Gujarat State

Abstract

Seasonal behavior of vascular cambium and development of secondary xylem were studied in the 2-3-years-old branches of Azadirachta indica A. Juss. (Meliaceae) growing in moist deciduous (MDF), dry deciduous (DDF) and scrub land (SF) forests of Gujarat State. Development of xylem began with the sprouting of new leaves in January in both MDF and DDF. Xylem production culminated in July in MDF whereas it reached peak in April and July in DDF. Cambial cells ceased to divide with the maturation of leaves in November and October in MDF and DDF, respectively. Mature xylem and phloem derivatives surrounded the cambium in December in MDF and November in DDF. Radial growth in the branches of trees growing in SF was found continuous throughout the year with peak cambial activity in April, July and November. In all the three forests, maximal radial growth was encountered in July when the rains were heavy. Cambial activity and xylem development showed significant correlation with the phenology of the trees in MDF and DDF whereas cambium was found active throughout the year and no correlation was found between the cambial activity and phenology in SF.     

IAA oxidase,peroxidase, and barrier zone formation in red maple

Wounding the cambium of red maple (Acer rubrum L.) altered peroxidase isoenzymes and enhanced the activities of peroxidase and indole-3-acetic acid oxidase in homogenates of adjacent cambial tissue. Four separate experiments were conducted, in May and July in each of two successive years. Wounds made in July caused greater increases in enzyme activity than those made in May. Alterations in peroxidase isoenzyme pattern were consistent for all experiments. These changes are intrepretable as aspects of the molecular regulation of barrier zone formation.