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1.
When the basal zones of 4-d-old hydroponically grown maize ( Zea mays L. cv. Seneca Horizon) roots were exposed to moist air for 2 d, the development of both endodermis and exodermis was affected. In the endodermis, Casparian bands enlarged and more cells developed suberin lamellae. The most striking effect was seen in the exodermis. In submerged controls, only 4% of the cells had Casparian bands, whereas in root regions exposed to air, 93% developed these structures. Similarly, in submerged roots 11% of the exodermal cells had either developing or mature suberin lamellae compared with 92% in the air-treated region. The majority of epidermal cells remained alive in the zone exposed to air. Some cell death had occurred earlier in the experiment when the seedlings were transferred from vermiculite to hydroponic culture. The precise stimulus(i) associated with the air treatment which led to accelerated development in both endodermis and exodermis is as yet unknown.  相似文献   

2.
Phialocephala fortinii Wang & Wilcox is a member of root-inhabiting fungi known collectively as dark septate endophytes (DSE). Although very common and distributed worldwide, few studies have documented their interaction with roots on a structural basis. The objective of this study was to determine the early colonization events and formation of microsclerotia of P. fortinii in roots of Asparagus officinalis L., a species known to have DSE. A loose network of hyphae accumulated at the root surface, and coils formed around root hairs and external to epidermal cells overlying short cells of the dimorphic, suberized exodermis. Root penetration occurred via swollen, appressorium-like structures into epidermal cells where coiling of hyphae occurred along the periphery of the cells. Hyphae penetrated from the epidermis into short exodermal cells and from these into cortical cells. Hyphae colonized the cortex up to the endodermis and sometimes entered the vascular cylinder. Some root tips were colonized as well. Microsclerotia in epidermal and exodermal short cells accumulated glycogen, protein, and polyphosphate. Energy-dispersive X-ray spectroscopy on distinct bodies visible in microsclerotial hyphae revealed high levels of phosphorus.  相似文献   

3.
The dimorphic exodermis of the root of onion (Allium cepa L.) consists of long and short cells, both of which have Casparian bands. The long cells and some of the short cells also have suberin lamellae. The proportion of short cells with lamellae increases with distance from the root tip and with plant age, but is not influenced by drought stress. In young regions of onion roots, characterized by a mature endodermis and an immature exodermis, the plasmalemma surface area that can be contacted by the soil solution is 90·9 mm2 per mm length of root, i.e. the sum of the plasmalemma surface areas of the epidermis, immature exodermis, cortical parenchyma and endodermis external to the Casparian band. This is reduced to 14·5–14·7 mm2 by the development of a Casparian band in the exodermis, which cuts off access to the cortical parenchyma, and by the development of suberin lamellae, which cut off access to the plasmalemmae of the long and some of the short cells of the exodermis. Death of all the epidermal cells, a consequence of drought, further reduces this area to 0·205–0·0183 mm2, i.e. the area of the outer tangential plasmalemmae of the short cells without suberin lamellae. In this condition, the root's capacity for ion uptake should be reduced but its capacity to resist water loss to the soil should be increased.  相似文献   

4.

Background and Aims

Most studies of exodermal structure and function have involved species with a uniseriate exodermis. To extend this work, the development and apoplastic permeability of Iris germanica roots with a multiseriate exodermis (MEX) were investigated. The effects of different growth conditions on MEX maturation were also tested. In addition, the exodermises of eight Iris species were observed to determine if their mature anatomy correlated with habitat.

Methods

Plants were grown in soil, hydroponics (with and without a humid air gap) or aeroponics. Roots were sectioned and stained with various dyes to detect MEX development from the root apical meristem, Casparian bands, suberin lamellae and tertiary wall thickenings. Apoplastic permeability was tested using dye (berberine) and ionic (ferric) tracers.

Key Results

The root apical meristem was open and MEX development non-uniform. In soil-grown roots, the exodermis started maturing (i.e. Casparian bands and suberin lamellae were deposited) 10 mm from the tip, and two layers had matured by 70 mm. In both hydro- and aeroponically grown roots, exodermal maturation was delayed. However, in areas of roots exposed to an air gap in the hydroponic system, MEX maturation was accelerated. In contrast, maturation of the endodermis was not influenced by the growth conditions. The mature MEX had an atypical Casparian band that was continuous around the root circumference. The MEX prevented the influx and efflux of berberine, but had variable resistance to ferric ions due to their toxic effects. Iris species living in well-drained soils developed a MEX, but species in water-saturated substrates had a uniseriate exodermis and aerenchyma.

Conclusions

MEX maturation was influenced by the roots'' growth medium. The MEX matures very close to the root tip in soil, but much further from the tip in hydro- and aeroponic culture. The air gap accelerated maturation of the second exodermal layer. In Iris, the type of exodermis was correlated with natural habitat suggesting that a MEX may be advantageous for drought tolerance.Key words: Iris germanica, roots, culture conditions, development, anatomy, apoplastic tracers, multiseriate exodermis, endodermis, root apical meristem  相似文献   

5.
Hair roots ofLysinema ciliatum R. Br. and some other Epacridaceaehave thick-walled cells in the epidermis. These are preferentiallycolonized with mycorrhizal fungi. Individual epidermal cellscontaining hyphal coils separate at the middle lamella and arereleased into the soil. Other colonized cells remain attachedto the roots, usually in groups, surrounded by bare exodermis,where epidermal cells have either collapsed or been sloughedoff. It is suggested that these colonized thick walled cellscan serve to prolong the mycorrhizal association and to infectnew hair roots as these emerge. The thick wall has a very specializedstructure and composition and could have a number of roles,either acting as a substrate or protective coat or in controllingwater status and uptake. Young hair-roots are surrounded bya mucilage sheath that is similar in appearance to that in Ericaceaeand apparently produced by root cap cells, not the epidermis. Lysinema ciliatum R. Br.; ericoid mycorrhiza; hair root; root cap; cortex; epidermis; exodermis  相似文献   

6.
Summary Onion (Allium cepa L. cv. Ebeneezer) roots from vermiculite culture were examined with transmission electron microscopy to detect the plasmodesmata in all tissues. In young root regions, plasmodesmata linked all living cells together in all directions. In old zones, the plasmodesmatal connections of the endodermis to its neighbor tissues were not interrupted by later suberin lamella and cellulosic wall deposition. Moreover, plasmodesmata in the fully mature endodermis usually exhibited a large central cavity. In the exodermis, however, upon deposition of suberin lamellae in long cells, all plasmodesmata that initially linked them to their adjacent cells were severed. Afterwards, the long cells lost the capability of forming wound pit callose and their protoplasts began to degenerate. The mature exodermal layer was symplastically bridged to its neighbors only by the short (passage) cells that lacked suberin lamellae. Compared to the long cells, the short cells not only had thicker cytoplasm surrounding their central vacuoles but also a higher density of mitochondria and rough endoplasmic reticulum, consistent with an active involvement in the transport processes of the root. The above results were obtained by an improved, extended transmission electron microscopy procedure devised to analyze plasmodesmata in cells with suberin lamellae. By prefixing root tissues in glutaraldehyde and acrolein, all cells were well preserved. Postfixation was carried out in osmium tetroxide at a low concentration (0.5%). Following dehydration in acetone and transfer to propylene oxide, infiltration with Spurr's resin was accomplished by incubating samples in the accelerator-free mixture for 4 days, then infiltrating samples in the accelerator-amended mixture for additional 4 days.Abbreviations IE immature exodermis - ME mature exodermis - TBO toluidine blue O - TEM transmission electron microscopy  相似文献   

7.
Hypoxia can stimulate the development of a suberized exodermis in aquatic plants; however, its influence on this aspect of terrestrial root development is sparsely documented. To determine the effects of hypoxia on maize (Zea mays cv. Seneca Horizon) roots, seedlings were grown in vermiculite (VERM), aerated hydroponics (AER), stagnant hydroponics with agar (STAG), or aerated hydroponics with agar (AERAG). The endo- and exodermis were examined for wall modifications. Lateral root emergence and aerenchyma formation were documented qualitatively. The endodermal Casparian band formation was unaffected by treatment. Endodermal and exodermal suberin lamella formation was earliest and most extensive in VERM. Suberization, especially in the exodermis of aerated treatments, was depressed in all hydroponic media. In comparison with AER, STAG exodermal lamellae were increased, but endodermal lamellae were decreased. Since the suberized exodermis forms a barrier to radial oxygen loss from roots to the medium, its stimulation in STAG roots (which also developed extensive aerenchyma) would help retain oxygen in the root. The reduction of endodermal lamellae should facilitate oxygen diffusion into the stele. Clearly, the response to environmental conditions is variable within individual cortical cell layers. Additionally, the observed patterns of lamellae, aerenchyma and lateral root development indicate a tight radial co-ordination of root development.  相似文献   

8.
The exodermis of Iris germanica roots is multiseriate. Its outermost layer matures first with typical Casparian bands and suberin lamellae. But as subsequent layers mature, the Casparian band extends into the tangential and anticlinal walls of their cells. Compared with roots in which the endodermis represents the major transport barrier, the multiseriate exodermis (MEX) was expected to reduce markedly radial water and solute transport. To test this idea, precocious maturation of the exodermis was induced with a humid air gap inside a hydroponic chamber. Hydraulic conductivity (Lp(pc)) was measured on completely submerged roots (with an immature exodermis) and on air-gap-exposed root regions (with two mature exodermal layers) using a pressure chamber. Compared with regions of roots with no mature exodermal layers, the mature MEX reduced Lp(pc) from 8.5×10(-8) to 3.9×10(-8) m s(-1) MPa(-1). Puncturing the MEX increased Lp(pc) to 19×10(-8) m s(-1) MPa(-1), indicating that this layer constituted a substantial hydraulic resistance within the root (75% of the total). Alternatively, a root pressure probe was used to produce pressure transients from which hydraulic conductivity was determined, but this device measured mainly flow through the endodermis in these wide-diameter roots. The permeability of roots to NaCl and ethanol was also reduced in the presence of two mature MEX layers. The data are discussed in terms of the validity of current root models and in terms of a potential role for I. germanica MEX during conditions of drought and salt stress.  相似文献   

9.
Patterns of infection withFusarium culmorum (W G Smith) Saccardo were observed in seedling roots of barley (Hordeum vulgare L.), wheat (Triticum aestivum L.), maize (Zea mays L.) and asparagus (Asparagus officinalis L). Apical regions of the main roots were not infected. Since penetration into the root occurred several days after inoculation and the roots were growing during the experiment, these regions had apparently not been in existence long enough to be infected. In older regions of barley, wheat and asparagus, hyphae entered through the tips of lateral roots. In barley and wheat, which had not developed any suberin lamellae in their subepidermal layer, infection occurred randomly over the remainder of the root. In maize, the fungus penetrated the epidermis at many sites but did not breach the exodermis in which all cells possessed both Casparian bands and suberin lamellae. Maize roots, therefore, sustained only minimal infections. In asparagus, the fungus grew through the short (passage) cells but never the long cells of the exodermis. In doing so, it penetrated cells possessing Casparian bands but lacking suberin lamellae. The results support the hypothesis that suberin lamellae provide effective barriers to the growth ofF. culmorum hyphae.  相似文献   

10.
Efficient rooting for establishment of papaya plantlets by micropropagation   总被引:2,自引:0,他引:2  
A low cost micropropagation protocol to produce high quality root systems which are easy and economical to acclimatize is essential for large-scale micropropagation of papaya (Carica papaya L.). In this study, individual shoots (>0.5 cm) with 23 leaves from in vitro papaya multiple shoots were cultured on MS agar medium containing 2.5 μM IBA under dark conditions for 1 week for root induction. They were then transferred to agar or vermiculite media, containing half strength MS medium, under aerated or non-aerated conditions, for root development. Rooting percentage of shoots cultured for 2 weeks in aerated vermiculite was 94.5%, compared with 90.0% in non-aerated vermiculite, 71.1% in aerated agar, and 62.2% in non-aerated agar. Shoots with roots were acclimated in vermiculite under 100% RH for 1 week and then under ambient conditions for 2 weeks in a temperature-controlled growth chamber (28 °C). The survival rates of the plantlets were 94.5% from aerated vermiculite, 87.8% from non-aerated vermiculite, 42.2% from aerated agar, and 35.6% from non-aerated agar. Thus, root induction in low-concentration IBA agar medium followed by root development in vermiculite containing half strength MS medium under aerated conditions results in efficient rooting of in vitro papaya shoots. This revised version was published online in June 2006 with corrections to the Cover Date.  相似文献   

11.
Water loss from roots back into drying soil is a problem ofpractical importance in plants growing under conditions of verylow substrate water potential, such as dry or saline areas.Root exodermis is relatively impermeable and has been suggestedto play a protective role against water loss. The relative waterretention ability was compared in root segments from exodermal(maize, onion, sunflower, Rhodes grass and sorghum) and non-exodermalspecies (Pisum sativum,Vicia fabaand wheat). Apical and basalsegments from exodermal roots, with different degrees of exodermisdevelopment, were also compared, as were segments from sorghumroots in which the exodermis thickness had been modified bysubjecting the plants to a 30 d water stress treatment. Waterretention was significantly higher in segments from exodermalroots. In each root, water loss was higher in apical than inbasal segments, regardless of the presence of exodermis. Insorghum, prolonged drought treatment increased exodermis thickeningin nodal roots, however, no differences in rates of water losswere observed in segments obtained from control and droughtedplants. Soil sheaths formed around roots of Rhodes grass growingin very dry soil with the epidermis adhering tightly to thesheath. In plants growing in the field, soil sheaths may bemore effective than the exodermis in preventing root water loss.Copyright1999 Annals of Botany Company. Root, exodermis, rhizosheaths, water loss.  相似文献   

12.
Gulnaz  A.  Iqbal  J.  Farooq  S.  Azam  F. 《Plant and Soil》1999,210(2):209-218
The paper investigates how the apoplastic route of ion transfer is affected by the outermost cortex cell layers of a primary root. Staining of hand-made cross sections with aniline blue in combination with berberine sulfate demonstrated the presence of casparian bands in the endo- and exodermis, potentially being responsible for hindering apoplastic ion movement. The use of the apoplastic dye Evan's Blue allowed viewing under a light microscope of potential sites of uncontrolled solute entry into the apoplast of the root cortex which mainly consisted of injured rhizodermis and/or exodermis cells. The distribution of the dye after staining was highly comparable to EDX analyses on freeze-dried cryosectioned roots. Here, we used Rb+ as a tracer for K+ in a short-time application on selected regions of intact roots from intact plants. After subsequent quench-freezing with liquid propane the distribution of K+ and Rb+ in cell walls was detected on freeze-dried cryosections by their specific X-rays resulting from the incident electrons in a SEM. All such attempts led to a single conclusion, namely, that the walls of the two outermost living cell sheaths of the cortex largely restrict passive solute movements into the apoplast. The ring of turgescent living rhizodermis cells in the root tip region forms the first barrier. With increasing distance to the root tip, in the course of their maturation resp. degradation, this particular function of the rhizodermis cells is replaced by the hypodermis resp. exodermis. Furthermore, the restriction of apoplastic ion flow by the outermost cortex cell layers is rather effective but not complete. Thus, the solute transfer into the stele is mainly restricted by the casparian bands of the endodermis. The overall conclusion is that the resistances of the rhizodermis and exodermis are additive to the endodermis in their role of regulating the apoplastic solute movement across roots. This revised version was published online in August 2006 with corrections to the Cover Date.  相似文献   

13.
Differential two-dimensional protein patterns as related to tissue specificity and water conditions were investigated within Brassica napus var oleifera root system. The different parts of the root system (tap root, lateral roots, and drought-induced short roots) were analyzed under various moisture regimes (regular watering at field capacity, progressive drought stress, and rewatering). Tissue specificity was evident from 25 differences in protein patterns (qualitative and quantitative) between well-watered lateral and tap roots. Twice as many polypeptides (52) were drought-affected and the response to the water stress was shown to be similar in both root types. In addition, more than half of the polypeptides detected as organ-specific were affected by drought. Based upon the trend of variation observed under drought and rehydration, three categories of polypeptides could be defined that might be differently involved in drought susceptibility or tolerance. A highly differentiated protein pattern characterized the drought-induced short roots. This pattern appeared as far from the watered as from the water-stressed normal roots. In particular, 13 unique polypeptides were detected which could be relevant to their adaptive morphogenesis and/or their specific drought tolerance induction. Upon rehydration, their polypeptide pattern and their specific morphology returned to a normal well-watered lateral root type.  相似文献   

14.
Abstract The development of suberin lamellae in the hypodermis of Zea mays cv. LG 11 was observed by electron microscopy and the presence of suberin inferred from autoliuorescence and by Sudan black B staining in nodal (adventitious) and primary (seminal) root axes. Suberin lamellae were evident at a distance of 30–50 mm from the tip of roots growing at 20°C and became more prominent with distance from the tip. Both oxygen deficiency and growth at 13°C produced shorter roots in which the hypodermis was suberized closer to the root tip. There were no suberin lamellae in epidermal cells or cortical collenchyma adjacent to the hypodermis. Plasmodesmata were not occluded by the suberin lamellae: there were twice as many of them in the inner tangential hypodermal wall (1,14 μn?2) as in the junction between the epidermis and hypodermis (0.54 μm?2). Water uptake by seminal axes (measured by micropotometry) was greater at distances more than 100 mm from the root lip than in the apical zone where the hypodermis was unsuberized. In the more mature zones of roots grown at 13°C rates of water uptake were greater than in roots grown at 20°C even though hypodermal suberization was more marked. Sleeves of epidermal/hypodermal cells (plus some accessory collenchyma) were isolated from the basal 60 mm of nodal axes by enzymatic digestion (drisclase). The roots were either kept totally immersed in culture solution or had the basal 50 mm exposed to moist air above the solution surface. In both treatments the permeabilities to tritiated water and 86Rb were low (circa 10?5mms?1) in sleeves isolated from the extreme base. In roots grown totally immersed, however, the permeability of sleeves increased 10 to 50-fold over a distance of 40 mm. In roots exposed to moist air the permeability remained at a low level until the point where the root entered the culture solution and then increased rapidly (> 50-fold in a distance of 8 mm). Growth of roots in oxygen depleted (5% O2) solutions promoted the development of extensive cortical aerenchymas. These developments were not associated with any reduction in permeability of sleeves isolated from the basal 40 mm of the axis. It was concluded that the presence of suberin lamellae in hypodermal walls does not necessarily indicate low permeability of cells or tissues to water or solutes. The properties of the walls (lamellae?) can be greatly changed by exposure to moist air, perhaps due to increased oxygen availability.  相似文献   

15.
The intensity of an ABA (abscisic acid) signal as a root-to-shoot signal, as well as its action on root hydraulic conductivity, strongly depends on the distribution of ABA during its radial transport across roots. Therefore ABA was visualized by immunolocalization with monoclonal ABA antibodies under conditions of lateral water flow induced by the application of a pressure gradient to the cut surface of the mesocotyl of maize seedlings. From the labelling of rhizodermis, hypodermis, cortical cells, and endodermis of roots of hydroponically (no exodermis) and aeroponically (with exodermis) grown seedlings it is concluded that the exodermis acts as a barrier to apoplastic transport that controls ABA uptake and efflux, but that the endodermis can easily be overcome via an apoplastic bypass. In longitudinal sections the strongest ABA signals originated from the root cap and the meristematic root tip, which is in agreement with the non-vacuolated cells of these tissues being an effective anion trap for ABA.  相似文献   

16.
All epidermal cells in root tips of panicoid grasses have been considered to be capable of hair formation. Observations made in this investigation suggested that cells of two maturation potentials may be present in the root-tip epidermis of Panicum virgatum. Protein bodies which swell and fuse in the region of elongation were revealed in the meristem of this grass by different staining procedures. In many roots not all cells seemed to receive the same amount of these bodies or of the protein-positive material which appeared to arise from them. Only deeply stained cells with large nucleoli were seen to form hairs. Epidermal cells of very hairy roots contained uniform nucleoli and exhibited similar distributions of protein material. The protein positive inclusions were never found in the cortex, a region of cells with one maturation potential. Following chloramphenicol treatment, root tips were found to contain epidermal cells with nucleoli of similar size, a reduced amount of protein bodies, and a reduction in the number of root hairs. RNase treatment did not appear to affect the integrity of the inclusions. The significance of such protein bodies is discussed in relation to differentiation of epidermal cells in P. virgatum.  相似文献   

17.
Cholewa E  Peterson CA 《Plant physiology》2004,134(4):1793-1802
The pathway of Ca(2+) movement from the soil solution into the stele of the root is not known with certainty despite a considerable body of literature on the subject. Does this ion cross an intact, mature exodermis and endodermis? If so, is its movement through these layers primarily apoplastic or symplastic? These questions were addressed using onion (Allium cepa) adventitious roots lacking laterals. Radioactive Ca(2+) applied to the root tip was not transported to the remainder of the plant, indicating that this ion cannot be supplied to the shoot through this region where the exodermis and endodermis are immature. A more mature zone, in which the endodermal Casparian band was present, delivered 2.67 nmol of Ca(2+) mm(-1) treated root length d(-1) to the transpiration stream, demonstrating that the ion had moved through an intact endodermis. Farther from the root tip, a third zone in which Casparian bands were present in the exodermis as well as the endodermis delivered 0.87 nmol Ca(2+) mm(-1) root length d(-1) to the transpiration stream, proving that the ion had moved through an unbroken exodermis. Compartmental elution analyses indicated that Ca(2+) had not diffused through the Casparian bands of the exodermis, and inhibitor studies using La(3+) and vanadate (VO(4)(3-)) pointed to a major involvement of the symplast in the radial transport of Ca(2+) through the endodermis. It was concluded that in onion roots, the radial movement of Ca(2+) through the exodermis and endodermis is primarily symplastic.  相似文献   

18.
Native bacteria, Pseudomonas and filamentous bacteria were quantified and localized on wheat roots grown in the field using fluorescence in situ hybridization (FISH). Seminal roots were sampled through the season from unploughed soil in a conservation farming system. Such soils are spatially heterogeneous, and many roots grow slowly through hard soil with cracks and pores containing dead roots remnant from previous crops. Root and rhizosphere morphology, and contact with soil particles were preserved, and autofluorescence was avoided by observing sections in the far-red with Cy5 and Cy5.5 fluorochromes. Spatial analyses showed that bacteria were embedded in a stable matrix (biofilm) within 11 microm of the root surface (range 2-30 microm) and were clustered on 40% of roots. Half the clusters co-located with axial grooves between epidermal cells, soil particles, cap cells or root hairs; the other half were not associated with visible features. Across all wheat roots, although variable, bacteria averaged 15.4 x 10(5) cells per mm(3) rhizosphere, and of these, Pseudomonas and filaments comprised 10% and 4%, respectively, with minor effects of sample time, and no effect of plant age. Root caps were most heavily colonized by bacteria along roots, and elongation zones least heavily colonized. Pseudomonas varied little with root development and were 17% of bacteria on the elongation zone. Filamentous bacteria were not found on the elongation zone. The most significant factor to rhizosphere populations along a wheat root, however, was contact with dead root remnants, where Pseudomonas were reduced but filaments increased to 57% of bacteria (P < 0.001). This corresponded with analyses of root remnants showing they were heavily colonized by bacteria, with 48% filaments (P < 0.001) and 1.4%Pseudomonas (P = 0.014). Efforts to manage rhizosphere bacteria for sustainable agricultural systems should continue to focus on root cap and mucilage chemistry, and remnant roots as sources of beneficial bacteria.  相似文献   

19.
以不同盐度下生长的互花米草(Spartina alterniflora)为材料,采用常规石蜡切片法对其根的横切结构进行显微观察,比较不同盐度下互花米草根结构的特点及变化规律,研究互花米草根对盐浸环境的适应性。观察结果显示:(1)互花米草根只有初生结构;(2)成熟根的表皮细胞基本毁坏、脱落;(3)互花米草根具有发达的外皮层和皮层通气组织,内皮层细胞壁五面加厚明显,且随盐度的升高呈先增大后减小的趋势;(4)维管柱中央被机械组织所填充,中柱鞘细胞壁也出现加厚现象。互花米草根的结构体现了其对盐浸环境的适应性特征。  相似文献   

20.
Rice plants were grown hydroponically and roots were prepared for light and electron microscopy using standard techniques. The roots are bounded by an epidermis, exodermis, and fibrous layer. The exodermis has a suberin lamella along its inner tangential wall. The fibrous layer is composed of thick-walled lignified cells with little pitting. The cortical parenchyma is compact when young, but expands and separates to form a zone of cell walls and air spaces in a spoked arrangement. Supporting columns of living parenchyma cells are occasionally present, particularly near lateral roots. The endodermis is typical for grasses with Casparian strips, suberin lamellae, and tertiary state walls with numerous pits. The pericycle and pith become sclerified. Protoxylem elements alternate with protophloem in the young root; later, early metaxylem, late metaxylem, and metaphloem proliferate. The exodermis, fibrous layer, lacunate cortex, and endodermis appear to present a formidable barrier to radial ion movement in the mature portions of the root.  相似文献   

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