DEVELOPMENT OF THE WOODY PORTION OF THE ROOT SYSTEM OF BETULA PAPYRIFERA

Our ontogenetic analysis of paper birch root systems shows that the fate of a root tip is related to its relative primary xylem diam (PXD). Lateral root tips with an initial PXD less than about 25 % of the parent root PXD are ephemeral. Some tips having a PXD of more than about 25 % of the parent root PXD become the permanent portions of the root system, enlarging over time as they elongate. In seedling root systems, the primary root and first-formed laterals are initially about the same size, and their PXD's all enlarge with increasing distance from the stem as the tips elongate to form the initial horizontal woody framework. Permanent lateral root branches with a large relative PXD develop after root tip injury, when the root tip is forced to grow in a curve, or from other unaccountable causes. Our observations show the importance of using relative diameters when classifying roots and when applying the concept of heterorhizy to paper birch root systems. Evidence points to the existence of some form of apical dominance in roots.     

勃氏甜龙竹不同部位根的解剖结构比较

该研究选择勃氏甜龙竹7月份(雨季)和11月份(干季)地面以下10 cm处土壤中秆基处生的根,秆下部节内(地上部分3~4节)和秆上部枝条基部在空气中自然生的根,以及7月份枝条高空压条产生的伸长阶段和成熟阶段的根作为实验材料,采用树脂切片和徒手切片法,对不同部位根的根尖和根毛区进行横切,甲苯胺蓝O和苏丹红7B染色,研究不同起源根的形态解剖结构,为竹子根解剖学研究提供新的理论信息。结果显示:(1)不同来源根的皮层宽度占根直径的比例、周缘纤维组织宽度及其细胞层数、内皮层细胞壁厚度均随生长时间延长而增加。(2)所有起源根的外皮层和内皮层细胞壁均有显著的木栓质沉积,但环内皮层木栓化程度存在显著差别。(3)测量不同来源根的后生木质部导管数目和直径发现,7月份采集的根导管数目更多、直径更大。(4)枝条基部自然生根的根尖与其他起源部位的根相比更为钝平,但根冠面积较小;秆下部节内自然生根非常硬,所以未能成功观察其根尖结构特点。研究认为,勃氏甜龙竹4种不同起源根的基本结构一致,但在各自的具体结构上还存在显著差别,可能与其生长的环境以及发育状况密切相关。     

Effect of Lateral Root Formation on the Vascular Pattern of Barley Roots

The vascular pattern in the root of barley (Hordeum vulgare L.), characterized by discontinuous xylem, is markedly affected by its branching. The roots become divided into unbranched segments alternating with branched segments with a more complex vascular pattern, formed by two systems differing in origin and age: the primary vascular system derived from the procambium and ontogenetically younger connective vascular system derived from stelar parenchyma. Adjacent to the sites of the lateral root initiation, reprogramming of parent stelar parenchyma for connective vascular elements occurs. The connecting phloem is represented by small sieve elements and companion cells, the connecting xylem is composed of small vessel elements with reticulate or scalariform-reticulate wall thickenings and simple perforations. Development of the connective vascular system secures continuous lateral and axial vascular connection between lateral root and parent root. The extent of the vascular connection in the parent root increases in an acropetal direction. Hydraulic effects of connective vascular tissue formation and parent root segmentation are discussed.     

DEVELOPMENT AND STRUCTURE OF THE VASCULAR CONNECTION BETWEEN THE PRIMARY AND SECONDARY ROOT OF GLYCINE MAX (L.) MERR.

The primary vascular connection between primary and secondary root of Glycine max (L.) Merr. was derived from stelar parenchyma and pericycle. Inner stelar parenchyma, associated with the parent metaxylem and outer stelar parenchyma adjacent to the pericycle, were resonsible for the histogenesis of the primary xylem connection. Acropetal maturation of the diarch xylem connection occurred after the lateral root emerged from the parent root. Development of tetrarchy occurred distal to the diarch xylem connection. The concentric primary phloem connection was derived from the pericycle and outer stelar parenchyma. Acropetal maturation of the primary phloem connection occurred prior to lateral root emergence from the parent root. Secondary growth quickly augmented the primary vascular connection. A substantial amount of mature secondary xylem formed prior to maturation of the secondary phloem. The structure of the primary and secondary vascular connections is described.     

ANATOMY OF SEEDLING ROOTS OF PSEUDOTSUGA MENZIESII

The seedling root system of Pseudotsuga menziesii (Mirb.) Franco consists of the primary root, active long laterals, long laterals that become mycorrhizal, and short roots that may or may not become mycorrhizal. Numerous adventitious roots arise from the pericycle in young roots and from the vascular cambium and pericycle in older roots following pruning. All actively growing apices have a single plate of initials, a complex zonation of mother cells, and a similar pattern of primary tissue differentiation. Short roots and mycorrhizal short roots have 2 plates of initials, one producing the stele and the other the root cap and cortex, and differentiation occurs close to the apex. Primary and adventitious roots are usually triarch, while long laterals are usually diarch as are all short roots. The latter lack secondary xylem, but mycorrhizal short roots may produce a small amount of secondary phloem.     

不同品种吊兰根初生木质部原型的观察与比较

以3个品种吊兰(金边吊兰、银心吊兰、银边吊兰)为材料,采用徒手切片的方法研究了吊兰根初生木质部原型与品种、生境及根直径的t关系。结果表明,不同品种的吊兰根初生木质部原型存在着差异;在水培条件下,吊兰根初生木质部原型与土培条件下的不同,水培条件下的根初生木质部束远少于土培条件;在水培条件下,吊兰根初生木质部原型与根的直径存在正相关关系,但是在土培条件下银心吊兰和银边吊兰根初生木质部原型则和根的直径没有关系。     

Water Flow Through Cotton Roots in Relation to Xylem Anatomy

The effect of root anatomy on water flow was studied in 7-d-oldcotton (Gossypium hirsutum L.) seedlings grown in solution culture.The total water flux of the intact root system was measuredusing a pressure chamber. Then successive terminal root sectionswere removed at 2,6,10 and 12 cm behind the root tip and theflux was remeasured after each successive cut was made. Xylemdevelopment at different distances behind the root apex wasstudied with a microscope using sections cut free-hand and stainedwith toluidine blue. Water flux increased with the removal ofsuccessive terminal root sections and this coincided with thedegree of basipetal primary xylem development. The large increasein water flux at 10 to 12 cm was associated with secondary xylemdevelopment and increased xylem vessel number. A comparison of water flow and xylem anatomy between roots withtetrarch (Stoneville 506 and Deltapine 41) and pentarch (T25strain) vascular bundle arrangements showed no significant differencesin the measured values of water flux for the primary root. Waterflux, estimated using Poiseuille's equation and measured xylemdimensions, was greater for the tetrarch roots, primarily becauseof the larger diameter of individual vessel elements. The increasednumber of vessel elements in the pentarch primary root of T25did not result in any apparent decrease in axial resistanceto water flow. Key words: Gossypium hirsutum L., roots, vascular bundle, xylem water flux, xylem     

Relative Contribution of Apices and Mature Tissues to ABA Synthesis in Droughted Maize Root Systems

We investigated whether different parts of maize root systemscould contribute to ABA synthesis, and whether a previous cycleof soil dehydration-rehydration would modify the ability ofroots to synthesize ABA. Maize (Zea mays L.) root tissues, i.e.mature primary root sections, young and unbranched primary rootsections, secondary roots and primary root tips, from both wellwatered plants and previously drought-rewatered plants, weresubjected to different degrees of dehydration and their ABAconcentration changes were assayed. All categories of rootsfrom always well watered plants, including mature tissues containingno apex, could synthesize ABA when dehydrated. Mature primaryroot sections and their previously associated secondary rootsaccumulated less ABA in response to dehydration than the youngprimary roots and primary root tips did, and their ABA accumulationwas not substantial until dehydration was below 65% of relativewater content (RWC). Previous soil dehydration-rehydration cyclessubstantially reduced ABA accumulation in these roots in responseto dehydration again. Young primary root sections and primaryroot tips accumulated ABA much more sensitively in responseto dehydration than mature root sections, although considerablevariations existed among different batches of young primaryroot sections. Results are discussed in the context of the relativecontribution of different categories of roots to ABA synthesiswhen the root system is in drying soil. We concluded that primaryroot apices should not contribute by more than 2% to the totalABA synthesis by the root system. (Received December 15, 1995; Accepted April 19, 1996)     

SOME ASPECTS OF VASCULAR DIFFERENTIATION IN ROOTS OF CASSIA

Hayat, Mohammed Arif (North Dakota State U., Fargo), and Charles Heimsch. Some aspects of vascular differentiation in roots of Cassia. Amer. Jour. Bot. 50(10): 965–971. Illus. 1963.—Vascular development, with emphasis on the differentiation of the protophloem, was studied in tips of primary roots of 18 species of Cassia. Variations in levels of protophloem sieve tube maturation were observed among roots of different species as well as among those of different length in the same species. In general, protophloem matured at greater distances from the apex in roots with the larger diameters. Compared with woody species, herbaceous species exhibited greater uniformity in levels of protophloem maturation, and this was correlated with greater uniformity in root diameter. Roots were either triarch or tetrarch. In some species with tetrarch roots, a change to a triarch pattern occurred during early growth. Structural changes in the differentiating root tip which involve the loss of a xylem arm and subsequent fusion of phloem strands are described.     

Tracheids in white spruce seedling's long lateral roots in response to nitrogen availability

This study examined how the availability of inorganic nitrogen (N) modified the anatomical characteristics of white spruce (Picea glauca (Moench) Voss) roots related to their hydraulic properties. Seedlings were grown for one growing season in 4 L capacity pots filled with sand under one of three N levels: low (10 ppm), medium (50 ppm) and high (125 ppm). First order lateral roots with intact tips were sampled from dormant seedlings in October. Root segments were collected from 4, 10, and 14 cm distances above the root tip for fixation and sectioning and for maceration. Additional specimens were collected from the 4 and 14 cm distances for maceration and scanning electron microscopy of xylem pits. Root diameter and surface area occupied by the xylem in root cross sections increased basipetally in all treatments but exceptions were found. Higher N-levels significantly increased root diameter and surface area occupied by the xylem. In the two higher N treatments secondary root development was more advanced near the root tip than in the low N treatment. There was a strong positive correlation between root diameter and cross-sectional root area occupied by the xylem (30–50% of the root cross section) but not in portions with little secondary development. Non-conducting space within the xylem occupied 10–13% of its cross-sectional surface. Tracheids of the primary xylem were larger, had larger lumens but thinner cell walls than those of the secondary xylem. Low N treatment seedling tracheids had smaller total cross-sectional area, less lumen, and less cell wall surface area than the two other N treatments. Tracheid diameter means were between 19–20 μm in the high and medium N treatments, and 15.2 μm in the low N treatment. The range was 4.5–51.3 μm. Tracheid length was not significantly affected by N. The average tracheid was about 1000 μm long, and the range was 110–3530 μm. Pit-border diameters ranged between 4.1–20.6 μm (average 10–11 μm) and were not affected by the N treatment. Pit aperture diameters were within 0.62–10.2 μm range (average between 3–4 μm) and were also not significantly affected by the N treatment, although tracheids from the medium N-treatment roots tended to have larger apertures. The pit border diameter equals that of the margo while the aperture size should be similar to that of the torus of the pit membrane. If the capacity for axial water transport in spruce roots is affected by N, it would be by its impact on conduit diameter and, possibly on the pit-membrane pore sizes but not by changes to conduit length and to the size of the pit membrane surface area. This revised version was published online in June 2006 with corrections to the Cover Date.     

栓皮栎根系解剖结构、水力特性及碳、氮含量研究

为阐明栓皮栎根系随径级的变化规律,探究其细根的合理划分标准。以1年生栓皮栎幼苗为研究对象,将其根系分为1、1～2、2～3、3～4 mm四个径级,分别制作石蜡切片观察解剖结构,比较木质部水力特性,测定碳氮含量及其比值,并采用主成分法对根系进行分类。结果表明:(1)随着径级增加,栓皮栎根系周皮、韧皮部和形成层组织厚度增加而占径比降低,木质部直径及其占径比均增加。(2)直径2 mm以上的栓皮栎根系木质部平均最大和最小导管直径、根比导水率和栓塞脆弱性指数增加显著; 而导管密度显著下降,导管面积与木质部面积之比变化不显著。(3)直径2 mm以上栓皮栎根系碳含量表现出显著增加,随着径级增加,根系氮含量下降、碳氮比升高。(4)主成分分析表明,13项根系结构和元素含量指标降维后,前2个主分量方差贡献率达62%,PCA双序轴显示栓皮栎根系可划分为2 mm以下的吸收根群和2 mm以上的运输根群。综上认为,以2 mm作为栓皮栎细根划分的标准兼顾了形态和功能的特点,更具有准确性。     

Effects of defoliation and symbiosis on polyamine levels in pine and birch

 We report the effect of ectomycorrhizal fungi (Suillus variegatus, Paxillus involutus) and defoliation on polyamine concentrations in pine (Pinus silvestris) and birch (Betula pendula) foliage and roots. Symbiotic root tips showed consistently higher concentrations of putrescine than non-symbiotic roots. Partial defoliation had no effect on the polyamine levels in mycorrhizal pine or birch roots. The foliage of mycorrhizal pine seedlings had lower putrescine concentrations and higher spermidine than foliage of non-mycorrhizal plants, and defoliation reversed this pattern. The response to partial defoliation differed in birch foliage: mycorrhizal status had no effect and all new growth after defoliation had higher spermidine levels than in non-defoliated birch. The potential role of polyamines in mycorrhizal symbiosis is discussed. Accepted: 26 February 1997     

Structural and Histochemical Features of the Slow-Growing Perennial <i>Coptis chinensis</i> Franch. (Ranunculaceae)

Huanglian (Coptis chinensis Franch.) is a slow-growing perennial medicinal herb with considerable economic value. This study aimed to determine the structural characteristics and the levels of berberine deposits in the organs and tissues of Huanglian using light and epifluorescence microscopy. The adventitious roots are composed of primary and secondary structures with endodermis, exodermis, and phellem. The rhizome structures are composed of primary and secondary structures with cuticle and phellem. The leaves are composed of sclerenchymatous rings, isolateral mesophyll, and thin cuticles. We detected berberine in the xylem walls of the roots and rhizomes as well as in the sclerenchymatous rings of the petioles. We postulate that as the exodermis is developed, the deposition of berberine in the xylem closest to the root tips may affect water and nutrient absorption and transfer. Leaf blades had a thin cuticle and isolateral mesophyll, suggesting shade tolerance. These structural and histochemical features suggest that Huanglian is adapted to the slow growing nature of a shady environment.     

VASCULAR CONNECTION BETWEEN LATERAL ROOTS AND STEM IN THE OPHIOGLOSSACEAE

The vascular connection between lateral roots and stem in the Ophioglossaceae and in two leptosporangiate fern species was examined. Two types of connections were found: “gradual” connections, which resemble leaf traces in ontogeny and morphology, and “abrupt” connections, which resemble the connections between lateral roots and their parent roots. Gradual root-stem connections occur in the genera Ophioglossum and Helminthostachys and in Woodwardia virginica. They are initiated in shoot apices distal to the level where cauline xylem elements mature. They resemble leaf traces in being provascular (procambial) strands that connect the cauline stele with the future vasculature of lateral appendages. As with leaf traces, gradual connections are part of the provascular and, later, protoxylem continuity between stems and lateral appendages. Gradual connections have many features in common with leaf traces, and the term root trace is applicable to them. The order of radial maturation of the primary xylem in gradual connections varies in different parts of the connections. It is endarch near the intersection with the cauline stele and exarch where the connections intersect root steles. Gradual connections resemble the transition regions of certain seed plants where protoxylem is also continuous from stem to root and the order of maturation is found to change continuously from stem to root. Abrupt connections occur in Botrychium and Osmunda cinnamomea. They develop in shoot apices at levels where cauline xylem is mature or maturing. The mature xylem does not dedifferentiate, so provascular and protoxylem continuity of the kind found in root traces does not occur. Also, reorientation of the order of maturation does not occur in abrupt connections. Xylem connectors are found in the region where radially oriented elements of the connections abut the longitudinally oriented cauline elements. Abrupt connections resemble the connection of secondary roots with their parent root systems since xylem connectors and the lack of continuity are also features found in these vascular systems. The resemblance of the vascular pattern of the fern root trace to the transition region of seed plants suggests that the radicle is more closely comparable to the cladogenous roots of pteridophytes than hitherto supposed.     

ANATOMY OF MACROZAMIA COMMUNIS LATERAL ROOTS AND ROOT NODULES FORMED IN VITRO STUDIED WITH LIGHT AND SCANNING ELECTRON MICROSCOPY

The anatomy of Macrozamia communis L. Johnson lateral roots and nodules was studied following axenic culture in light and darkness. Pointed lateral roots from dark cultures had an open apical organization similar to that of other cycads and gymnosperms. A distinct protoderm-derived epidermis was not observed. At the apex, the dermis was formed by the outer root capcortical cell layer. Subapically, the outer cortex formed the dermis. No evidence of an algal zone was observed in these roots. The stele was bounded by a distinct endodermis and contained an exarch, diarch xylem. Apogeotropic nodules which developed at the root-shoot junction in darkness, branched dichotomously and had rounded tips covered by tangentially-enlarged root cap cells. The root cap was reduced to a few cell layers and was confined to the extreme nodule apex. The central region of the apical meristem was enlarged, and meristematic cells contained differentiated amyloplasts. A presumptive algal zone was present in some but not all nodules and divided the cortex into inner and outer regions. Stelar anatomy was similar to that observed in pointed, dark-grown lateral roots, except that there was greater xylem differentiation. Nodules which developed in the light were similar to dark-formed nodules, except that root cap cells were radially enlarged and extended over the flanks of the nodule forming a persistent root cap. The heteromorphic lateral roots of M. communis formed a developmental continuum not a heterorhizic root system.     

侧柏扦插不定根发生模式研究

该研究以侧柏一年生硬枝插穗为实验材料,利用连续组织切片技术观察插穗不定根发生发育过程中的组织结构变化,分析插穗外部形态变化、不定根原基起源和不定根的形成过程,探讨侧柏插穗不定根发生模式和不定根的组织学起源。结果显示:侧柏扦插后可由愈伤组织、皮部诱导产生不定根,出现皮部生根、愈伤组织生根、愈伤组织兼具皮部生根3种类型;侧柏插穗中存在少量潜伏根原基,但插穗生根类型以诱导生根为主;不定根原基诱导产生于愈伤组织、木质部、形成层及次生韧皮部等部位。研究认为侧柏扦插生根属于多位点发生模式,不定根原基的组织学起源是愈伤组织、髓射线、射线原始细胞、尚未分化成熟的木质部细胞,通过人工诱导同时激活这些不定根起源位点能够显著提高生根率和生根质量。     

Investigation of the infection process of macrophomina phaseolina on the surface of soybean roots using scanning electron microscopy

Sclerotial germination, hyphal growth pattern, and appressorial formation of the charcoal rot fungus, Macrophomina phaseolina (Tassi) Goid. on the surface of soybean roots, Glycine max (L.) Merrill was observed using scanning electron microscopy. Sclerotial germination is followed by a hyphal growth pattern which does not appear to be associated with any specific structural feature on the corrugated root surface. Appressoria are produced on the root surface at the tips of both primary hyphae and side branches within three days after inoculation.     

Stem hypoxia and root respiration of flooded maple and birch seedlings

Some researchers have attributed flood tolerance of woody plants to air entering the shoot through stems, leaves, or lenticels and diffusing to the roots to sustain aerobie respiration. The purpose of this study was to determine if internal aeration of roots by lower stems, changes in gross morphology of lower stems, or both, contribute to flood tolerance of certain tree species. Greenhouse-grown seedlings of red maple ( Acer rubrum L.) and river birch ( Betula nigra L.) tolerated at least 30 days of flooding, where as sugar maple ( Acer saccharum Marsh) and European white birch (also called silver birch, Betula pendula Roth) were intolerant. Flood treatment induced lentieel intumescences and adventitious root formation on red maple stems, but only adventitious roots formed on river birch stems. Stem morphology of sugar maple and European birch was unchanged by flooding. Flood stress decreased oxygen consumption capacity of excised roots from both tolerant and intolerant species. Exclusion of oxygen from the lower stems of flooded red maple and river birch prevented lenticel intumescence and adventitious root formation, but flood tolerance and root respiration capacity were unchanged. Neither internal aeration nor changes in stem morphology appear to account for flood tolerance of red maple and river birch.     

Mycorrhizas and C and N transformations in the rhizospheres of Pinus sylvestris,Picea abies and Betula pendula seedlings

Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) seedlings were grown in a greenhouse for four months in three different soils. The soils were from a field afforestation site on former agricultural land: soil from a pine site, soil from a spruce site and soil from a birch site. Pots without seedlings were included. The aim was to discover, independent of the effects of the different quality of aboveground litter and microclimate under the tree species, whether the roots change the microbial activities and chemical characteristics of the soil, whether the changes are dependent on the tree species, and whether the changes vary in different soils. Pine, spruce and birch had, on average, five, one and six meters of roots, respectively. Birch had by far the highest number of short root tips, on average 11 450 per seedling, compared to 1900 and 450 in pine and spruce seedlings, respectively. The majority of the short roots of pine and spruce were brown sheathed mycorrhizas, and those of birch were mycorrhizas in an early stage of development. The seedlings caused no major changes in either the soil pH or the concentrations of nutrients in the soils, but did affect the microbial characteristics of the soils. The effect of the tree species did not differ in different soils. Microbial biomass C and N, C mineralization rate and the concentration of ergosterol were all higher under birch and pine than under spruce and in plantless soils. Nitrate concentrations were lowest under pine and birch, but rates of net N mineralization, nitrification and denitrification did not differ under different seedlings. The stimulative effect of pine and especially birch on soil microbes was possibly due to them having more roots and releasing more root exudates to soil. There were, however, indications that not only the length/mass of roots determined the changes in microbial activities, but also differences in root activities per unit of root or in the quality of root exudates.     

Water relations in silver birch during springtime: How is sap pressurised?

• Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known.
• We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period.
• The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations.
• Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed.