Location of the low temperature water flow barrier in stems

Experiments are described indicating the magnitude and location of the low temperature barrier to lateral water flow in stems of cotton (Gossypium hirsutum L. `Auburn 7-683'). Rehydration of wilted stem tissues was performed at 6 C and 32 C. Compared with the 32 C control, a 13-fold increase in the rehydration halftime was recorded at 6 C when water entered the secondary phloem tissues across the vascular cambium from the secondary xylem. However, only a 3-fold increase in the rehydration halftime occurred when water entered phloem tissues through the cortex, and most of this increase was due to the higher viscosity of water at the lower temperature. These results show that the cambial region of an intact cotton stem markedly resists the radial flow of water at lower temperatures. This resistance was not demonstated by other stem tissues.     

Oxygen isotope enrichment of organic matter in Ricinus communis during the diel course and as affected by assimilate transport

The oxygen isotope composition in leaf water and organic compounds in different plant tissues is useful for assessing the physiological performance of plants in their environment, but more information is needed on Delta(18)O variation during a diel course. Here, we assessed Delta(18)O of the organic matter in leaves, phloem and xylem in stem segments, and fine roots of Ricinus communis during a full diel cycle. Enrichment of newly assimilated organic matter in equilibrium with leaf water was calculated by applying a nonsteady-state evaporative enrichment model. During the light period, Delta(18)O of the water soluble organic matter pool in leaves and phloem could be explained by a 27 per thousand enrichment compared with leaf water enrichment. Leaf water enrichment influenced Delta(18)O of phloem organic matter during the night via daytime starch synthesis and night-time starch remobilization. Phloem transport did not affect Delta(18)O of phloem organic matter. Diel variation in Delta(18)O in organic matter pools can be modeled, and oxygen isotopic information is not biased during transport through the plant. These findings will aid field studies that characterize environmental influences on plant water balance using Delta(18)O in phloem organic matter or tree rings.     

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

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

白鲜营养器官解剖结构及其与白鲜碱的积累关系

应用植物解剖学、组织化学及植物化学方法对白鲜营养器官根、茎、叶的结构及其生物碱的积累进行了研究。结果显示:(1)白鲜根的次生结构以及茎和叶的结构类似一般双子叶植物;白鲜多年生根主要由周皮、次生韧皮部、维管形成层以及次生木质部组成,根次生韧皮部中可见大量的淀粉、草酸钙簇晶、韧皮纤维以及油细胞;茎由表皮、皮层、维管组织和髓组成;叶由表皮、栅栏组织、海绵组织和叶脉组成;在茎和叶初生韧皮部的位置均分布有韧皮纤维,在叶表皮上分布有头状腺毛和非腺毛;在茎和叶紧贴表皮处分布有分泌囊。(2)组织化学分析结果显示:在白鲜多年生根中,生物碱类物质主要分布在周皮、次生韧皮部、维管形成层和木薄壁细胞中;在茎中,生物碱主要分布在表皮、皮层、韧皮部、木薄壁细胞及髓周围薄壁细胞中;在叶中,生物碱主要分布在表皮细胞、叶肉组织和维管组织的薄壁细胞;此外在分泌囊和头状腺毛中亦含有生物碱类物质。(3)植物化学结果显示,秦岭产白鲜根皮/白鲜皮、根木质部、茎和叶中白鲜碱含量分别为0.041%、0.012%、0.004%和0.002%,其中木质部中白鲜碱含量和其他部分地区白鲜皮中白鲜碱含量类似。研究表明,在秦岭产白鲜营养器官中,除根皮/白鲜皮外,在根木质部亦含有大量的白鲜碱,且在茎和叶中亦含有一定的白鲜碱,具有潜在的开发利用价值。     

SUCCESSIVE CAMBIA IN THE STEM OF PHYTOLACCA DIOICA

Phytolacca dioica L., an evergreen tree of the Phytolaccaceae, is one of the species of Phytolacca which shows anomalous secondary thickening in its stem. This mode of thickening has been regarded as successive cambial activity or alternatively, in some more recent interpretations, as thickening by unidirectional activity of a cambial zone. The stem thickening of P. dioica is of the former type. The cambium produces fascicular strands, showing centrifugal differentiation of xylem and centripetal differentiation of phloem on opposite sides of the cambial layer, and rays are produced between the fascicular areas. In both xylem and phloem the younger elements are closer to the cambium than the older elements. Succeeding cambia arise periodically by periclinal divisions in a layer of parenchyma cells two or three cells beyond the outermost intact phloem derived from the current cambium. Each cambium forms a few parenchyma cells on both sides before it forms derivatives which mature into lignified xylem elements or conductive elements of the phloem. The parenchyma thus formed toward the outside later becomes the site of the origin of the succeeding cambium. Only one or two layers of this phloem parenchyma go on to form the new cambium; the remaining cells accumulate between the outermost phloem and the cortex. P. weberbaueri shows stem structure similar to P. dioica. P. meziana, a shrub, shows normal stem structure.     

Stem anatomy of Dolichos lablab Linn (Fabaceae): Origin of cambium and reverse orientation of vascular bundles

Secondary growth in the stem of Dolichos lablab is achieved by the formation of eccentric successive rings of vascular bundles. The stem is composed of parenchymatous ground tissue and xylem and phloem confined to portions of small cambial segments. However, development of new cambial segments can be observed from the obliterating ray parenchyma, the outermost phloem parenchyma and the secondary cortical parenchyma. Initially cambium develops as small segments, which latter become joined to form a complete cylinder of vascular cambium. Each cambial ring is functionally divided into two distinct regions. The one segment of cambium produces thick-walled lignified xylem derivatives in centripetal direction and phloem elements centrifugally. The other segment produces only thin-walled parenchyma on both xylem and phloem side. In mature stems, some of the axial parenchyma embedded deep inside the xylem acquires meristematic activity and leads to the formation of thick-walled xylem derivatives centrifugally and phloem elements centripetally. The secondary xylem comprises vessel elements, tracheids, fibres and axial parenchyma. Rays are uni-multiseriate in the region of cambium that produces xylem and phloem derivatives, while in some of the regions of cambium large multiseriate, compound, aggregate and polycentric rays can be noticed.     

Distribution of an Indoleacetic Acid-oxidase-inhibitor in the Storage Root of Daucus carota

Indoleacetic acid (IAA)-oxidase from both secondary phloem and xylem was dependent on 2,4-dichlorophenol for activity, and was enhanced by addition of Mn²⁺. The pH optimum was 6.0 from both tissues. IAA-oxidase and its inhibitors were distributed differently in the secondary phloem and secondary xylem of carrot root. In the phloem a high IAA-oxidase activity was distributed uniformly along the radius but in the xylem a somewhat lower concentration decreased from the cambium. IAA-oxidase inhibitor in the phloem increased exponentially from a very low concentration near the cambium, whereas in the xylem an appreciable concentration was present near the cambium, decreasing linearly with distance from the cambium. Longitudinal gradients in the xylem parallel studies by other workers with the greatest IAA-destroying capacity present in older tissues. In the xylem inhibitor decreased and IAA-oxidase increased from the root apex. In the phloem IAA-oxidase was uniform, whereas the inhibitor increased in older tissue.

The IAA-oxidase inhibitors in phloem and xylem may be different. In the xylem the IAA-oxidase inhibitor may be a lignin precursor present in young cells which disappears as lignification proceeds. In the phloem IAA-oxidase reacting with endogenous IAA appears to form a physiologically active product.

     

Radial secondary growth and formation of successive cambia and their products in Ipomoea hederifolia L. (Convolvulaceae)

Ipomoea hederifolia stems increase in thickness using a combination of different types of cambial variant, such as the discontinuous concentric rings of cambia, the development of included phloem, the reverse orientation of discontinuous cambial segments, the internal phloem, the formation of secondary xylem and phloem from the internal cambium, and differentiation of cork in the pith. After primary growth, the first ring of cambium arises between the external primary phloem and primary xylem, producing secondary phloem centrifugally and secondary xylem centripetally. The stem becomes lobed, flat, undulating, or irregular in shape as a result of the formation of both discontinuous and continuous concentric rings of cambia. As the formation of secondary xylem is greater in one region than in another, this results in the formation of a grooved stem. Successive cambia formed after the first ring are of two distinct functional types: (1) functionally normal successive cambia that divide to form secondary xylem centripetally and secondary phloem centrifugally, like other dicotyledons that show successive rings, and (2) abnormal cambia with reverse orientation. The former type of successive rings originates from the parenchyma cells located outside the phloem produced by previous cambium. The latter type of cambium develops from the conjunctive tissue located at the base of the secondary xylem formed by functionally normal cambia. This cambium is functionally inverted, producing secondary xylem centrifugally and secondary phloem centripetally. In later secondary growth, xylem parenchyma situated deep inside the secondary xylem undergoes de‐differentiation, and re‐differentiates into included phloem islands in secondary xylem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 30–40.     

牛膝茎的发育解剖学研究

应用植物解剖学方法研究了牛膝茎的发育过程。研究结果表明,牛膝茎的发育包括原分生组织、初生分生组织、初生结构、次生结构和三生生长5个发育阶段。原分生组织具有典型分生组织的细胞特征;初生分生组织包括原表皮、基本分生组织和原形成层。在茎的发育过程中,初生生长和早期的次生生长是正常的,但在次生生长过程中,次生维管组织仅有束中形成层产生,而没有束间形成层的分化和活动。茎的三生生长是由维管柱外侧保留的原形成层细胞发生的额外形成层的活动产生的。额外形成层开始只向内交替产生三生木质部和其间的结合组织,后来向外产生三生韧皮部,形成一轮三生维管束。牛膝茎内的韧皮纤维来源于原形成层,应属于原生韧皮部性质。牛膝茎中的2个外韧型髓维管束也来源于原形成层,与正常维管束在位置上没有相关性。但其结构类型具有多样性,有时可形成不完全的周木型髓维管束。     

Comparative Studies on Differentiation of Regenerated Tissue in Broussonetia papyrifera

This paper describes the differentiation process of regenerated tissue after ordinary girdling or after removal of a section of xylem from the stem, and the disparity in differentiation of the regenerated tissues after being differently treateds in Broussonetia papyrifera. After ordinary girdling for 3–4 weeks, new bark regenerated in the xylem. During the process of rind' formation, many specks of meristematic tissue were formed in the callus, from which vascular tissue clusters were developed. In addition, the new periderm appeared almost at the same time as the new vascular cambium was seen. When a section of xylem was removed from the stem, numerous calli developed rapidly on the inner surface of the bark. Meanwhile, the vascular cambium appeared in the immature phloem. Soon after, discontinued meristematic tissue bands also occurred in the callus. These meristematic tissues then connected with each other to form a concave oblate cambial ring which developed xylem inward and phloem outward. About 2–3 weeks later, the concave oblate trunk grew lengthwisely connecting with the upper anct lower portions of the normal stem. By then, the tree continued to grow. The inner surface tissue of the bark, after the xylem was removed, differentiated about one week earlier than the tissue on the surface of the xylem after girdling.     

Contrasting dynamics of water and mineral nutrients in stems shown by stable isotope tracers and cryo‐SIMS

Lateral exchange of water and nutrients between xylem and surrounding tissues helps to de‐couple uptake from utilization in all parts of a plant. We studied the dynamics of these exchanges, using stable isotope tracers for water (H₂¹⁸O), magnesium (²⁶Mg), potassium (⁴¹K) and calcium (⁴⁴Ca) delivered via a cut stem for various periods to the transpiration stream of bean shoots (Phaseolus vulgaris cv. Fardenlosa Shiny). Tracers were subsequently mapped in stem cross‐sections with cryo‐secondary ion mass spectrometry. The water tracer equilibrated within minutes across the entire cross‐section. In contrast, the nutrient tracers showed a very heterogeneous exchange between xylem vessels and the different stem tissues, even after 4 h. Dynamics of nutrients in the tissues revealed a fast and extensive exchange of nutrients in the xylem parenchyma, with, for example, calcium being completely replaced by tracer in less than 5 min. Dilution of potassium tracer during its 30 s transit in xylem sap through the stem showed that potassium concentration was up‐regulated over many hours, to the extent that some of it was probably supplied by phloem recirculation from the shoot.     

Periodicity of Cambial Activity and Changes of Starch Content in Broussonetia papyrifera

Methods of sampling and sections preparaction were the same as reported previously. Except that sampling was made at monthly intervals between May 20 and July 30, then at 7–14 day-intervals between July 30 and October 14, and then at monthly intervals between October 14 and March 25 in the next year. The stored starch in various tissues was stained with PAS reaction. During active period of cambium in Broussonetia papyrifera after July 30, the cell layers of immature xylem and phloem decreased progressively, and the formation of mature xylem and phloem increased rapidly. The formation of late wood started early in August, formation of xylem ceased after September 5, followed by ceasation of phloem formation about 1.5 months later. Increasing and decreasing of stored starch were closely related to the periodicity of cambial activity during the year. Starch grains decreased progressively after cambial activity was resumed in early spring until they disappeared in all the stem tissues. Then, starch accumulated progressively again after cambial activity slowed down, particularly after the ceasation of xylem formation. However, after the formation of phloem had ceased, the stored starch once again disappeared progressively until the end of December, and accumulated again. Such changes might be related to the transition of cambium activity involving two periods of dormancy.     

Changes of Peroxidase and Estease Isozymes During Periodicity of Cambial Activity in Broussonetia papyrifera (L.) Vent.

Methods of sampling and sections preparaction were the same as reported previously. Except that sampling was made at monthly intervals between May 20 and July 30, then at 7–14 day-intervals between July 30 and October 14, and then at monthly intervals between October 14 and March 25 in the next year. The stored starch in various tissues was stained with PAS reaction. During active period of cambium in Broussonetia papyrifera after July 30, the cell layers of immature xylem and phloem decreased progressively, and the formation of mature xylem and phloem increased rapidly. The formation of late wood started early in August, formation of xylem ceased after September 5, followed by ceasation of phloem formation about 1.5 months later. Increasing and decreasing of stored starch were closely related to the periodicity of cambial activity during the year. Starch grains decreased progressively after cambial activity was resumed in early spring until they disappeared in all the stem tissues. Then, starch accumulated progressively again after cambial activity slowed down, particularly after the ceasation of xylem formation. However, after the formation of phloem had ceased, the stored starch once again disappeared progressively until the end of December, and accumulated again. Such changes might be related to the transition of cambium activity involving two periods of dormancy.     

构树形成层的活动周期及其淀粉贮量的变化

在构树（Ｂｒｏｕｓｓｏｎｅｔｉａ ｐａｐｙｒｉｆｅｒａ （Ｌ．） Ｖｅｎｔ．）形成层活动周期中,每年７月末以后,未成熟的木质部和韧皮部逐渐减少,成熟的木质部和韧皮部急剧增多。８月初开始分化晚材。进入９月后木质部的形成逐渐停止,而一个半月以后才停止形成韧皮部。淀粉贮量的消长与形成层的活动周期有很强的相关关系。早春形成层恢复活动后,淀粉贮量逐渐减少直至消失。尔后,形成层活动减慢,特别是木质部分化停止后,淀粉又开始积累。当韧皮部分化也停止后,淀粉又消失,直至翌年１月才重新积累,这似乎与两个休眠期的转化有关     

Diurnal pattern of water potential in woody plants

The dynamic relationship between the rates of water loss and uptake controls plant water status. Marked diurnal variations in water potential of both leaves and fruit occurred in all plants studied. Variations in water status during the day were most clearly related to changes in evaporative demand of the air and were different for the east and west sides of a tree. At night, the plant water potential reflected the soil moisture status.

Changes in the water potential of pear fruit were correlated with changes in fruit diameter. Since water loss from fruit occurred mostly through the pedicel into the xylem of the tree, the fruit could be used as a crude gauge of xylem water potential, which also showed dramatic changes during the day.

     

Water relations of tropical dry forest flowers: pathways for water entry and the role of extracellular polysaccharides

Many trees in tropical dry forests flower during the dry season when evaporative demand is high and soil water levels are low. In this study the factors influencing the water balance of flowers from three species of dry forest trees were examined. Flowers had greater mucilage contents than leaves, high intrinsic and absolute capacitances, long time constants for water exchange and high transfer resistances. Flower water potentials were higher than in leaves and did not fluctuate over the lifespan of the flower. Flower water content also remained constant even though evaporation rates were high, suggesting that water was being supplied from the stem. In two of the species, the water potential gradient between flowers and leaves was opposite to that necessary for water transport from stem to flowers through the xylem, and it was therefore hypothesized that water may enter the flower through the phloem. Calculations showed that nectar production in these flowers could drive a sink of sufficient magnitude to allow water input via the phloem equal to water lost from the flower to the atmosphere.     

白皮松形成层的活动周期及其多糖贮量和淀粉酶同工酶的变化

4月初,白皮松(Pinus bungeana Zucc.)形成层带细胞开始增大,未成熟的木质部和韧皮部细胞增多,下旬出现成熟的木质部细胞。5月以后,木质部和韧皮部的形成速度加快,6月初进入晚材形成期。8月初停止产生木质部,9月中旬停止产生韧皮部。多糖颗粒的消长与形成层活动有较强的相关性,恢复活动前后颗粒含量持续增长,6月进入晚材形成期才持续减少,至翌年1月初完全消失,3月又重新积累,并迅速达到高峰。淀粉酶同工酶在活动期只有一条酶带,形成层停止产生木质部后出现了3条特异酶带,12月初又出现了2条特异酶带,这5条酶带都一直存在到形成层恢复活动。     

Tissue-Specific Expression of Cell Wall Proteins in Developing Soybean Tissues

Cell wall hydroxyproline-rich glycoproteins (HRGPs) and glycine-rich proteins (GRPs) were examined at the protein and at the mRNA levels in developing soybean tissues by tissue print immunoblots and RNA blots. In young soybean stems, HRGPs are expressed most heavily in cambium cells, in a few layers of cortex cells surrounding primary phloem, and in some parenchyma cells around the primary xylem, whereas GRPs are highly expressed in the primary xylem and also in the primary phloem. In older soybean stems, HRGP genes are expressed exclusively in cambium cells and GRP genes are most heavily expressed in newly differentiated secondary xylem cells. Similar expression patterns of HRGPs and of GRPs were found in soybean petioles, seedcoats, and young hypocotyls, and also in bean petioles and stems. HRGPs and GRPs become insolubilized in soybean stem cell walls. Three major HRGP mRNAs and two major GRP mRNAs accumulate in soybean stems. Soluble HRGPs are abundant in young hypocotyl apical regions and young root apical regions, whereas in hypocotyl and root mature regions, soluble HRGPs are found only in a few layers of cortex cells surrounding the vascular bundles. GRPs are specifically localized in primary xylem cell walls of young root. These results show that the gene expression of HRGPs and GRPs is developmentally regulated in a tissue-specific manner. In soybean tissues, HRGPs are most heavily expressed in meristematic cells and in some of those cells that may be under stress, whereas GRPs are expressed in all cells that are or are going to be lignified.     

风花菜根异常次生结构的解剖学观察

利用光镜观察了风花菜（Rorippa islandica (Oed.)Borb.）的根的发育过程中的解剖结构的变化,发现风花菜根具有异常次生结构,第一年早期次生结构中,在导管周围形成额外形成层并由额外形成层向内外分裂形成大量的薄壁组织,从而使根快速增粗,在根的横切面上,导管分散在薄壁组织当中。第一年晚期的次生结构中,由木质部中薄壁组织细胞反分化形成额外形成层,许多分散的额外形成层片段连接形成形成层环,形成层环向内向外分裂形成三生木质部及三生韧皮部,从而构成三生维管组织环,三生维管组织环形成方向由外向内,可以形成多环。第二年风花菜的韧皮部以及皮层中的薄壁组织反分化形成额外形成层,并且由额外形成层片段连接形成额外形成层环,进而分裂形成三生维管组织环,而且在皮层中可以形成多层,形成的方向为由内向外。风花菜根不同发育时期的异常次生结构是与其生理活动相适应的。