Certain seasonal changes in balsam fir cambium and its derivatives

Summary The seasonal cambial activity of balsam fir producing normal and compression wood was observed. Samples were taken throughout one growing season. Activity was initiated in normal wood with the precocious development of one sieve cell per file 2 weeks before activity was initiated in compression wood with the formation of one new sieve cell per file. Deactivation dates in normal and compression wood were the same.Dating of the phloem adjacent to normal wood can be carried out using crushed sieve cells or an annually formed band of tannin filled parenchyma whereas adjacent to compression wood, only crushed sieve cells can be used.The maximum cambial zone widths in normal and compression wood were 15 and 16 cells, respectively. The final xylem to phloem ratio in normal wood was 14:1 while in compression wood it was 21:1.Time required for development of tracheids was examined throughout the growing season. Growth trends for the radial growth and differentiation phases were studied in both normal and compression wood. Tracheid development in normal wood took between 14 and 33 days while in compression wood it varied between 5 and 49 days.Research financed under McIntire-Stennis Research Project 5009 and COE Projects R625-173 and R625-195     

Growth, cambial activity and phloem structure in compatible and incompatible peach/plum grafts

Peach (Prunus persica L. Batsch) shoots grafted onto compatible or incompatible clones of rootstocks of myrobolan plum (Prunus cerasifera L. Ehrh. cv. myrobolan) were observed in a greenhouse for 100 days after grafting. The incompatible grafts showed foliar symptoms of incompatibility and reduced shoot growth about 60 days after grafting. Light microscopic studies revealed that cambial activity stopped earlier in the rootstocks, than in the scions, of incompatible grafts. Structural modifications were also observed in the phloem of incompatible grafts. Compared with the compatible graft rootstock, the number of sieve elements differentiated from the cambial zone was reduced in the incompatible graft rootstock, whereas the production of parenchyma cells was not affected. No important ultrastructural alteration was observed in the sieve tubes of the incompatible grafts. However, osmiophilic granulations near the plasma membrane of sieve plates were observed more frequently in the rootstocks of incompatible grafts than in ungrafted myrobolan controls. It is concluded that the external symptoms of incompatibility are not related to massive structural modifications or degeneration of the conducting tissues.     

Regular cambial activity and xylem and phloem formation in locally heated and cooled stem portions of Norway spruce

The effect of heating (23–25°C) and cooling (9–11°C) on regular cambial activity and xylem and phloem formation in the stem portion of Norway spruce was investigated. Adult trees were sampled at 21-day intervals during the 2005 vegetation period. Continuously elevated temperatures increased the rate of cell division in the first part of the growing season, but did not significantly prolong cambial activity at the end of the vegetation period in the heated tree. Low-temperature treatments shortened regular cambial activity and slowed down cell production. The xylem growth ring was wider in the heated sample and narrower in the cooled sample compared to the control. The temperature in the cambial region was only negligibly transferred along the stem from the site of its application. In general, the temperature in the cambium was affected by a long-term rise or drop in air temperatures. Both experiments affected the structure and width of phloem growth increments. The tangential band of the axial parenchyma was not continuous in the cooled sample. The number of late phloem cells was reduced in the cool-treated sample and increased in the heat-treated sample. Our experiments confirmed the effect of constantly increased or decreased temperatures on regular cambial activity in Norway spruce.     

Understanding the role of the cytoskeleton in wood formation in angiosperm trees: hybrid aspen (Populus tremula x P. tremuloides) as the model species

The involvement of microfilaments and microtubules in the development of the radial and axial components of secondary xylem (wood) in hybrid aspen (Populus tremula L. x P. tremuloides Michx.) was studied by indirect immunofluorescent localization techniques. In addition to cambial cells, the differentiated cell types considered were early- and late-wood vessel elements, axial parenchyma, normal-wood fibers and gelatinous fibers, and contact and isolation ray cells. Microfilaments were rare in ray cambial cells, but were abundant and axially arranged in their derivatives once cell elongation had begun, and persisted in that orientation in mature ray cells. Microfilaments were axially arranged in fusiform cambial cells and persisted in that orientation in all xylem derivatives of those cells. Microtubules were randomly oriented in ray and fusiform cells of the cambial zone. Dense arrays of parallel-aligned microtubules were oriented near axially in the developing gelatinous fibers, but at a wide range of angles in normal-wood fibers. Ellipses of microfilaments were associated with pit development in fiber cells and isolation ray cells. Rings of co-localized microtubules and microfilaments were associated with developing inter-vessel bordered pits and vessel-contact ray cell contact pits, and, in the case of bordered pits, these rings decreased in diameter as the over-arching pit border increased in size. Although only microtubules were seen at the periphery of the perforation plate of vessel elements, a prominent meshwork of microfilaments overlaid the perforation plate itself. A consensus view of the roles of the cytoskeleton during wood formation in angiosperm trees is presented.     

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。     

油桐带化病中发现的类细菌（BLO）

油桐(Aleurites fordii Hemsl.)是陕西省紫阳县重要的经济林树种之一,但在该县发现油桐感染带化病严重,病害呈点片状发生,根据在洞河、城关等地区调查,一般发病率为3％,严重的达7％,发病的油桐树龄为2—5年,以3年生幼树发病较严重。油桐带化病病变形态与香椿带化病病变十分相似。感病的油桐树在未发生病变的前一年,树高有生长迅速的特点,一般在病枝以下的一节(于两侧枝之间)一年内高生长达3—4m。病树最显著的病状是新生的枝条呈扁平的带状,尖端卷曲或扭曲生长,如同一把大刀,有的形成齐头铲状,最宽处可达13cm,病枝叶片明显变小(图5);当枝梢孕育花芽亦能开花座果,果实仅有正常成熟果实大小的1/4,萎缩枯死,影响油桐树的正常生长及桐油产量。此病的蔓延阻碍了油桐树的发展。     

Anatomical and cellular responses of Pinus monticola stem tissues to invasion by Cronartium ribicola

White pine blister rust (Cronartium ribicola) causes extensive damage to white pines and their associated ecosystems across North America. The anatomical and cellular characteristics of C. ribicola colonization in Pinus monticola branch and stem tissues were studied as a basis for understanding host tree reactions that may be related to resistance. Samples examined showed typical fusiform swelling and some had produced aecia. The reaction of phloem and xylem tissues was compared with non-infected tissue using light and electron microscopy. Cortical parenchyma and phloem polyphenolic parenchyma cells underwent mitotic division, cell swelling, and ca sixfold greater accumulation of phenolic compounds in colonized vs. control stems. In the cortex and secondary phloem, haustoria were common in parenchymatous cells, and hyphae were abundant in the intercellular spaces, but cell death was rare, unless aecia had ruptured the stem cortex. Hyphae were also common in xylem rays, tracheids and between tracheids. Disease-induced changes in the cambial zone included development of cambium-derived xylem traumatic resin ducts. Results demonstrate that diverse host defence responses were activated in the bark of apparently susceptible trees, but lack of mechanical damage by C. ribicola to the phenol-containing host cells and the resin duct system allowed extensive colonization and development of aecia despite elicitation of these stem defences. Interactions between P. monticola and C. ribicola are discussed and compared with other conifer–fungus pathosystems.     

果树果实细胞超微结构研究进展

从细胞超微结构综述了仁果类、核果类、浆果类果树果实的薄壁细胞、筛管伴胞（SE／CC）复合体、韧皮薄壁细胞及各种细胞器的研究进展,并对果实细胞超微结构的发展方向提出了自己的看法和建议。     

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.     

Anatomical and FTIR analyses of phloem and xylem of <Emphasis Type="Italic">Tetracentron sinense</Emphasis>

The fast growth of Tetracentron sinense is a potential valuable timber resource, but whether its anatomy and chemical components are suitable for timber is unknown. We used light microscopy and SEM to examine the anatomical structure and FITR to measure the chemical components of the phloem and xylem of this tree. Radial variations in growth ring width and tracheid dimensions were also evaluated. The sieve tube, phloem parenchyma cell and sclereids clusters were the main cells in phloem, and the tracheid was the fundamental cell in xylem. An unusual tracheid type, fiber-tracheids or vessel-liked elements was visible. Wood rays nonstoried, uniseriate and multiseriate, including heterogeneous II, occasionally I, and usually 3–6 cells wide. The mean growth-ring width was 2.53 ± 0.46 mm, and the percentage of late wood was over 60%. For radial variation, growth-ring width increased at an early growth stage, and reached the largest increment during years 11–15, then decreased. The maximum growth-ring width was 5.313 mm. During late growth (60–85 years), trees also maintained a high radial growth increment. Radial variation in the percentage of late wood was uniform, about 50–70%, throughout the growth years. Growth patterns in the length and width of early and late wood were similar as the trees aged. From the FTIR results, the chemical components differed significantly between xylem and phloem, hemicellulose in particular was higher in the xylem than in the phloem, where it was apparently absent. All of these suggest that the composition of phloem in T. sinense is very similar to that of hardwood, and it has higher growth ratio and uniform wood properties.     

Definitive Callose Deposition in Tapping Panel Dryness Affected Bark of Hevea brasiliensis

The structural and functional aspects of sieve elements in Tapping Panel Dryness (TPD) affected bark of Hevea brasiliensis, with special reference to definitive callose deposition, have been studied using histochemical methods. The effect of ethephon in definitive callose deposition in TPD affected bark also has been investigated, as the ethylene is reported to have a major role in TPD. The phloem of Hevea is characterized by sieve elements with compound sieve plates. The deposition of definitive callose on the sieve areas following TPD limits the translocation of sap through sieve elements. The amount of functional phloem was narrow and the accumulation of definitive callose was more in the sieve elements of TPD affected bark compared to that of unaffected bark of the same tree and of healthy trees. The affected area showed an increment in deposition of definitive callose in both vertical and horizontal extent of sieve areas; and there was a reduction in length of sieve plate, density of sieve pores, length of inter-sieve area wall, and number of sieve element rows in the inner bark. The lateral walls of non-functional sieve elements became thick and lignified. The anatomical changes in the sieve elements of TPD affected bark suggest that the translocation of phloem sap is regulated by deposition of definitive callose and less differentiation of sieve elements from vascular cambium. The external application of ethephon on bark enhanced the definitive callose deposition on sieve elements. The findings from the study reveal that ethylene plays a major role in the senescence of sieve elements.     

Potassium distribution in black heartwood of sugi (Cryptomeria japonica) I: Localization in axial parenchyma cells

This is the first report to ascertain potassium location in black heartwood of sugi (Cryptomeria japonica). The objective of this study was to understand the distribution of potassium in sugi black heartwood in connection with anatomical morphology. Scanning electron microscopy with energy dispersive X-ray analysis (SEM-EDXA) was used to investigate the distribution of inorganic elements, especially potassium. In black heartwood, potassium was detected in all tissues (tracheids, ray parenchyma, and axial parenchyma), but its concentrations were different in each xylem tissue. Potassium was particularly accumulated in the droplets in axial parenchyma cells. In addition, calcium was also detected in all tissues. Phosphorus sometimes existed in the granules in ray parenchyma cells. From optical microscopy, the axial parenchyma cells were observed to be arranged in a tangential direction on transverse section and the droplets in these cells were distributed in areas of high ash and green moisture content. For red heartwood, potassium detected by SEM-EDXA was far lower in all tissues than in the black heartwood, while the trend for calcium was the converse. Potassium in axial parenchyma cells was more abundant in black heartwood than in typical pale red heartwood.     

Role of periderm in resistance of Eucalyptus marginata roots against Phytophthora cinnamomi

Phytophthora cinnamomi Rands invaded phloem tissue preferentially in Eucalyptus marginata Sm. (Jarrah) roots of both primary and secondary structure. Rate of fungal growth in the “primary” roots was less than in roots with secondary thickening. In the main field study jarrah roots between 1.5–2.5 mm in diameter, were inoculated with P. cinnamomi at regular intervals, over 12 months and in most cases invasion of the phloem was halted within the first 8 weeks of infection. In the majority of roots which exhibited secondary structure, lesion shape reflected the pattern of periderm development. Some lesions were peripheral in the outer phloem as cylindrical periderms had formed close to the vascular cambium. Other lesions were limited to a sector of the roots; periderm had formed radially through the secondary phloem and a zone of suberization was often continuous through the cambial zone and xylem. Formation of periderm at the edge of lesions was a multi-step process with various changes occurring, including lignification and suberization of walls and accumulation of autofluorescent compounds in cell walls and vacuoles. The layers of suberized and lignified cells formed near lesion fronts, were necrophylactic periderms. Necrophylactic periderm formation generally preceded the establishment of periderm of the exophylactic type.     

Can the structure of dormant cambium and the widths of phloem and xylem increments be used as indicators for tree vitality?

We investigated the structure and width of the dormant cambium and of the increments of phloem and xylem of Quercus robur to estimate their potential as indicators for tree vitality. The samples were taken from three woodlands, two in Slovenia [Krakovo forest (KRA) and Murska Suma (MUS)] and one in Croatia [Kobiljak (KOB)], with reported tree decline. The number of dormant cells seems to reflect the initial capacity of the cambium to accomplish cell division. With the exception of two trees at KRA, cell production was always higher on the xylem side than on the phloem side. The annual phloem increments were narrower, less variable among trees and with clear lower and upper limits. With increased cambial cell productivity, the share of the xylem in the total annual radial increment increased following a curvilinear function. In trees with an annual radial increment >3.5 mm, the xylem size represented more than 90 % of the total radial growth. The anatomical variables analyzed show that the most limiting environmental conditions seem to prevail at KRA, whereas the conditions at MUS seem to be most favorable in terms of radial growth. Analysis of the width and structure of xylem and phloem increments, the number of dormant cambial cells and their inter-relationships can provide additional information on the vitality of oaks.     

漆树科4种植物次生韧皮部的解剖比较

【目的】以漆树科3属4个种(盐肤木、青麸杨、黄连木和黄栌)的次生韧皮部为研究对象,阐明各物种次生韧皮部组成细胞的结构特征和分布规律,为植物次生韧皮部组成细胞结构特征与分泌结构之间的关系研究提供理论依据,并为漆树科植物系统演化的相关研究提供参考依据。【方法】采集盐肤木、青麸杨、黄连木和黄栌的树皮样品(样品含形成层、次生韧皮部及周皮,不含次生木质部),采用比较解剖学的方法,对漆树科4种植物的次生韧皮部的结构特征、组成细胞的构成及其排列方式进行比较,并对4种植物的乳汁道结构进行研究。【结果】漆树科4种植物次生韧皮部均是由筛管分子和伴胞、韧皮薄壁组织细胞、射线细胞、乳汁道和韧皮纤维构成,呈切向带状相间排列。黄连木、青麸杨和盐肤木的具功能韧皮部和无功能韧皮部分界明显;青麸杨的射线细胞在无功能韧皮部中倾斜分布,其余3种植物均为整齐纵向排列;青麸杨的乳汁道为分枝型乳汁道,其他3个种的乳汁道均无分枝,仅有少量相邻乳汁道融合现象发生;黄连木和黄栌次生韧皮部内分布有大量的韧皮纤维,青麸杨韧皮部中较少,而盐肤木韧皮部中无韧皮纤维分布。【结论】漆树科4种植物次生韧皮部组成细胞及乳汁道结构存在一定差异,青麸杨乳汁道是目前为止漆树科植物中发现的较为稀有的分枝型乳汁道,研究结果能够为漆树科系统进化研究提供参考依据。     

Palm “wood”

The fine structure of differentiating and mature sieve elements, tracheary elements and fibers of palms has been studied. The fine structure of sieve elements in general resembles that of the sieve elements in angiosperms except that not all the species of palms have P-protein. Material examined after taking precautions to minimize sudden release of pressure in the phloem indicate that the sieve-plate pores may be free of occlusions in intact sieve elements. Sieve elements in the basal parts of the stem estimated to be several decades old maintain their structural integrity and apparently remain functional throughout the life of the palm. Differentiating xylem elements are characterized by the presence of amoeboid plastids with electron-dense granular stroma. All the organelles normally present in meristematic cells are also present in differentiating tracheary elements. Bundles of microfilaments are often present in such elements. Structural changes that occur as the tracheary element matures are comparable to those observed in tracheary elements of other angiosperms. Xylem parenchymatous cells do not have the so called protective wall layer. The most striking ultrastructural feature in differentiating fibers of the root is the deep invagination of plasmalemma that traverses almost the entire width of the cell. The secondary wall of the fibers is characteristically multilayered.This paper was presented at the Eighth Cellulose Conference, Syracuse, New York, U.S.A.     

Induction of anatomically based defense responses in stems of diverse conifers by methyl jasmonate: a phylogenetic perspective

Conifers have evolved constitutive and inducible defense mechanisms to help in both wound healing and defense against attack by bark beetles and other organisms. These defenses include oleoresin, phenolics, and static structures in secondary phloem, such as lignified cells and calcium oxalate crystals, that create physical barriers. We used a phylogenetic approach to investigate the defense anatomy of conifer stems of 13 species from five families following treatment with methyl jasmonate (MJ), a compound that induces defense responses in stems of several Pinaceae species. Methyl jasmonate induced a response similar to wounding except that the response was not accompanied by lesion formation, necrosis or a hypersensitive response. In the Pinaceae species studied, MJ induced polyphenolic parenchyma (PP) cell activation and xylem traumatic resin duct (TD) formation. Members of the Taxodiaceae, which are not known to produce large quantities of resin, showed massive xylem TD formation and surface resinosis following MJ treatment. Treatment with MJ caused members of the Araucariaceae and Cupressaceae to form axial phloem resin ducts but not xylem ducts, whereas Podocarpaceae species showed no induction of resin-producing structures. All species treated with MJ showed phenolic deposition in PP cells, and early lignification of phloem fibers was observed in most of the non-Pinaceae species. We conclude that, although evolution of resin-producing structures occurred independently in conifer lineages, MJ seems to induce resin production regardless of tissue location, as well as inducing deposition of phenolic compounds. Co-evolution of conifer defensive strategies and bark beetle pests is discussed.     

Variation of cell length in bark and wood of tropical trees

Summary The variation in length of cell elements both in bark and wood of a few tropical trees (Mangifera altissima, Mansonia altissima, Pentacme contorta, Pterocarpus marsupium, Shorea megrosensis, Shorea squamata, Triplochtion scleroxylon) is described. The phloem fibres and sieve tube members show a tendency for increase in length from periderm to cambium, just as the wood fibres and vessel members increase from pith to cambium. The phloem fibres are generally longer than wood fibres (40–70%). In woods with storeyed cambia the sieve tube members exhibit no variation, retaining a constant length throughout the width of the bark; the phloem fibres possess the normal trend of a length increase from periderm to cambium. The length variation of sieve tube members and phloem fibres with the height of the tree generally exhibits a decreasing trend from butt to top, although a slight increase in length of phloem fibres at the top was noticed.Presented at the IUFRO Division V Meeting, September 1973, Cape Town, South AfricaThanks are due to Mrs. K. Puchstein for help with the measurements. The financial support by the Deutsche Forschungsgemeinschaft and the Gesellschaft der Freunde und Förderer der Bundesforschungsanstalt für Forst- und Holzwirtschaft, Reinbek, is also gratefully acknowledged.