Studies onCabomba aquatica (Cabombaceae)

Cabomba aquatica has been found for the first time in India. Both, submerged and floating leaves are epistomatic. The stomata are anomocytic and perigenous in their development. The metaxylem vessel elements have simple perforation plates and spiral side wall thickening. Vessel elements are found in roots, rhizomes and aerial stems. The venation pattern is of two types like the dimorphic leaves. On the basis of these and other featuresCabomba deserves family rank.     

SEM studies on vessels in ferns. 11. Ophioglossum

With scanning electron microscopy (SEM), the nature of metaxylem vessel elements and tracheids was examined in Ophioglossum crotalophomides, 0. pendulum subsp. falcatum , and 0. vulgatum roots and rhizomes. Vessels were identified in all species. End walls of vessel elements, which bear perforations, are like lateral wall pitting of those elements in the secondary wall framework and differ only in absence of pit membranes or presence of pit membrane remnants. Some of the perforations contain pit membrane remnants that have large pores, small porosities, or are threadlike or weblike in structure. Dimorphic perforations were found in some vessel elements of rhizomes of 0. pendulum subsp. falcatum. Tracheids are very likely present in addition to vessels in all three species. The secondary wall framework of both tracheids and vessels is basically scalariform, although deviations in pattern are present. Vessel elements of Ophiglossum are entirely comparable to those of leptosporangiate ferns.     

PRESENCE OF VESSELS IN WOOD OF SARCANDRA (CHLORANTHACEAE); COMMENTS ON VESSEL ORIGINS IN ANGIOSPERMS

Sarcandra is the only genus of Chloranthaceae hitherto thought to be vesselless. Study of liquid-preserved material of S. glabra revealed that in root secondary xylem some tracheary elements are wider in diameter and have markedly scalariform end walls combined with circular pits on lateral walls. Examination of these wider tracheary elements with scanning electron microscope (SEM) demonstrated various degrees of pit membrane absence in the end walls. Commonly a few threadlike fibrils traverse the pits (perforations); these as well as intact nature of pit membranes in pits at ends of some perforation plates are evidence that lack of pit membranes does not result from damage during processing. Some perforations lack any remnants of pit membranes. Although perforation plates and therefore vessels are present in Sarcandra roots, no perforations were observed in tracheary elements of stems or lignotubers. Further, stem tracheids do not have the prominently scalariform end walls that the vessel elements in roots do. Presence of vessels in Sarcandra removes at least one (probably several) hypothetical events of vessel origin that must be postulated to account for known patterns of vessel distribution in angiosperms, assuming that they are primitively vesselless. Seven (perhaps fewer) vessel origin events in angiosperms could account for these patterns; two of those events (Nelumbo and monocotyledons) are different from the others in nature. Widely accepted data on trends of vessel specialization in woody dicotyledons yield an unappreciated implication: vessel specialization has happened in a highly polyphyletic manner in dicotyledons, and therefore multiple vessel origins represent a logical extension backward in time. If a group of vesselless dictyoledons ancestral to other angiosperms existed, they can be hypothesized to have had a relatively homogeneous floral plan now that Sarcandra-like plants no longer need be imagined within that group. Sarcandra and other Chloranthaceae show that the borderline between vessel absence and presence is less sharp than generally appreciated.     

云南箭竹不同年龄段假鞭的解剖学结构特征及木质化观察

该研究以云南箭竹不同年龄段的假鞭为实验材料,采用滑动切片法并利用光学显微镜观察,分析云南箭竹假鞭的解剖结构特征及其随年龄的动态变化,为假鞭结构研究提供新的解剖学数据信息。结果显示:(1)云南箭竹假鞭节间的表皮层只有1层细胞,皮下层由3～4层细胞壁加厚的纤维细胞组成,皮层一般有20～25层不规则的薄壁细胞,成熟的皮层细胞会形成皮层气道,髓实心不具髓腔。(2)云南箭竹假鞭纤维壁厚随鞭龄增加而增加,且同一年龄假鞭的内侧韧皮部面积大于外侧;纤维腔径随鞭龄增加而逐渐减小,但同一年龄假鞭内侧纤维腔径大于外侧;韧皮部的面积、维管束和导管的直径均随着鞭龄的增加而增大。(3)假鞭维管束一般不具有原生导管,外部维管通常有2个较大的后生导管,在假鞭中部及内部通常只有1个后生导管,另1个后生导管不发育或发育不全。(4)在0.5年生到2年生的云南箭竹假鞭中,被染成紫红色的木质素在纤维细胞壁、薄壁细胞壁、导管细胞壁中都有分布,且随着假鞭年龄的增加染色逐渐加深,表明云南箭竹假鞭木质素含量随着鞭龄的增长而不断增加,木质化程度随鞭龄的增长逐渐提高。     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE. V. DOES NELUMBO HAVE VESSELS?

The examination of the primary xylem of roots of Nelumbo has revealed the presence of vessel elements with intra-face, transversely-rowed to obliquely-rowed bordered pitted side walls and scalariformly perforated end walls. Whole mounts of macerated xylem, serial sections of roots, and cells through which a water suspension of India ink particles under slight or no pressure was passed, were the means of investigation. It is our opinion that this discovery does not support theories which derive monocotyledons from the Nymphaeaceae. A discussion leading to this conclusion is presented.     

Accumulation of high levels of potassium in the developing xylem elements in roots of soybean and some other dicotyledons

Summary Potassium concentrations have been determined by cryo-analytical scanning microscopy in vacuoles of cells of the roots of soybean and six other dicotyledons. Developing vessel elements accumulate the highest concentration of potassium in any cell type in these roots, and those of the secondary xylem have more (median 190 mM normalized to 120%) than those of either the early (median 100 mM, normalized 80%) or late metaxylem (median 110 mM, normalized 100%). Potassium concentration in these developing vessels always exceeds those of their adjacent parenchyma (60–80%), which in turn are higher than those in cells of the cortex (30–40%), including the endodermis. Potassium concentration in the vessels increases during their development until cell death and maturation when it drops dramatically (normalized 4–12%). Developing vessel elements are clearly major sites of potassium accumulation in roots and need to be considered in any models of ion uptake, accumulation, circulation, or exudation.Abbreviations EDX energy dispersive X-ray microanalysis - EMX early metaxylem - K elemental potassium - LMX late metaxylem - CSEM cryo-scanning electron microscopy - SX secondary xylem Dedicated to the memory of Professor John G. Torrey     

Solute concentrations in xylem sap along vessels of maize primary roots at high root pressure

The elemental composition of xylem sap has been determined by cryo-analytical microscopy in situ along vessels in the roots of maize plants frozen intact while root pressure was high. The only chemical element (including carbon) present in significant concentrations in the vessels was potassium at 11 mM and 15 mM in the late (LMX) and early (EMX) metaxylem, respectively. There was no gradient of [K] along the vessels, which each run the length of the mature proximal end of the roots. At the distal end of each vessel, in the oldest still living vessel, which each run the length of the mature proximal end of the roots. At the distal end of each vessel, in the oldest still living vessel elements, there was sharp rise in [K] to 110 mM and 130 mM in the LMX and EMX, respectively.     

Nelumbonaceae:Systematic position and species diversification revealed by the complete chloroplast genome

<正>Nelumbonaceae is a morphologically unique family of angiosperms and was traditionally placed in Nymphaeales;more recently,it was placed in Proteales based on molecular data,or in an order of its own,Nelumbonales. To determine the systematic position of the family and to date the divergence time of the family and the divergence time of its two intercontinentally disjunct species,we sequenced the entire chloroplast genome of Nelumbo lutea and most of the chloroplast genes of,N.nucifera.We carried out phylogenetic and molecular dating analyses of the two species and representatives of 47 other plant families,representing the major lineages of angiosperms, using 83 plastid genes.The N.lutea genome was 163 510 bp long,with a total of 130 coding genes and an overall GC content of 38%.No significant structural differences among the genomes of N.lutea,Nymphaea alba, and Platanus occidentalis were observed.The phylogenetic relationships based on the 83 plastid genes revealed a close relationship between Nelumbonaceae and Platanaceae.The divergence times were estimated to be 109 Ma between the two families and 1.5 Ma between the two Nelumbo species.The estimated time was only slightly longer than the age of known Nelumbo fossils,suggesting morphological stasis within Nelumbonaceae.We conclude that Nelumbonaceae holds a position in or close to Proteales.We further conclude that the two species of Nelumbo diverged recently from a common ancestor and do not represent ancient relicts on different continents.     

SEM studies on vessels in ferns. 2. Pteridium

Xylem from roots and rhizomes of two infraspecific taxa of Pteridium aquilinum was studied by means of scanning electron microscopy (SEM). All tracheary elements proved to be vessels. End wall perforation plates were all scalariform, lacked pit membrane remnants in at least the central part of the perforation plate, and varied with respect to width of bars, from wide to tenuous, and with respect to presence of pit membrane remnants. In addition, porose pit membranes on walls that are likely all lateral vessel-to-vessel walls must be considered to be perforations also, although different from those on end walls. Lateral wall perforation plates, hypothesized by one worker on the basis of tylosis presence but denied by another on the basis of light microscopy, were confirmed by demonstration of pores with SEM. In addition, lateral walls of Pteridium vessels bear some grooves interconnecting pit apertures; this feature is newly figured by SEM for ferns. Lateral wall pitting that is not porose may either have striate thickenings of the primary wall or be smooth. Vessel presence and degree of specialization in Pteridium vessels may bear a relationship to the wide ecological tolerances of the genus.     

Fossil Nelumbonaceae from the La Colonia Formation (Campanian–Maastrichtian,Upper Cretaceous), Chubut,Patagonia, Argentina

We describe here fossil leaves and fruits assignable to the family Nelumbonaceae Dumortier 1828. Fossils were collected at the Cañadón del Irupé locality, La Colonia Formation (Campanian–Maastrichtian, Upper Cretaceous), Chubut Province, Patagonia, Argentina. The fossil leaves are simple, with symmetrical and peltate lamina, orbicular in shape, and an entire margin. The venation is truly actinodromous with at least 15 primary veins, the secondary veins are poorly developed and intercalated with the primary veins, the third category vein is opposite, and the areoles are well-developed and 4- to 5-sided. These features are characteristic of the extant genus Nelumbo Adamson 1763, and the presence of these characters in the fossils allows their placement within this genus. Reproductive structures, similar to the fruit-receptacle of Nelumbo with fruits in situ, were found associated but not in organic connection with the leaves. These fossils constitute the only and oldest record for the family in the Southern Hemisphere. This record indicates that Nelumbo was more widespread in the past than today.     

Late Maturation of Large Metaxylem Vessels in Soybean Roots: Significance for Water and Nutrient Supply to the Shoot

The large, late metaxylem (LMX) in the roots of soybean beginsdevelopment in the centre of the stele after lignification ofthe early metaxylem poles. Subsequent maturation of the firstappearing LMX elements is gradual. They were never mature inthe 8-d-old seedlings examined. In 10 to 15-d-old plants thefirst LMX matured to open vessels at a mean of 17 cm proximalto the root tip. Additional LMX vessels developed in more proximalregions of the roots and these also matured gradually. Based on calculations from relative vessel diameters, the potentialflow of xylem sap in a single central LMX vessel is 50 timesthat in the total of all the early metaxylem (EMX) vessels ofa typical primary root of soybean. There was a marked dependence of relative leaf area on the lengthof primary root with open LMX vessels. This may result fromthe predicted increased water and nutrient flow to the shoot,facilitated by the opening of the large vessels. It is suggestedthat, as in maize, the living LMX elements may function in ionaccumulation. Dicotyledonous roots, soybean, Glycine max, xylem vessels, xylem maturation, water conduction     

Root xylem embolisms and refilling. Relation To water potentials of soil, roots, and leaves, and osmotic potentials of root xylem Sap

Embolism and refilling of vessels was monitored directly by cryomicroscopy of field-grown corn (Zea mays L.) roots. To test the reliability of an earlier study showing embolism refilling in roots at negative leaf water potentials, embolisms were counted, and root water potentials (Psiroot) and osmotic potentials of exuded xylem sap from the same roots were measured by isopiestic psychrometry. All vessels were full at dawn (Psiroot -0.1 MPa). Embolisms were first seen in late metaxylem vessels at 8 AM. Embolized late metaxylem vessels peaked at 50% at 10 AM (Psiroot -0.1 MPa), fell to 44% by 12 PM (Psiroot -0.23 MPa), then dropped steadily to zero by early evening (Psiroot -0.28 MPa). Transpiration was highest (8.5 μg cm-2 s-1) between 12 and 2 PM when the percentage of vessels embolized was falling. Embolized vessels were refilled by liquid moving through their lateral walls. Xylem sap was very low in solutes. The mechanism of vessel refilling, when Psiroot is negative, requires further investigation. Daily embolism and refilling in roots of well-watered plants is a normal occurrence and may be a component of an important hydraulic signaling mechanism between roots and shoots.     

Anatomy of Seedling Roots of Tropical Maize (Zea mays L.) Cultivars at Low Water Supply

Establishment of maize seedlings can be difficult at low soilmoisture content. Anatomy of root metaxylem vessels may influencethe capacity for water transport and respective genotypic differencesmight be useful for selection purposes. To test this, six tropicalmaize (Zea mays L.) cultivars were grown in large PVC tubescontaining a sandy substrate at 5% (M5) and 10% (M10) moisturecontents for 2 weeks. The percentage changes in root diametersdue to M5 was similar for most cultivars but differed for mainroot types. Root diameters were not consistently related tometaxylem structure, but in a few cases, thin roots had smallerdiameter metaxylem vessels. The M5 treatment reduced the numberof late metaxylem vessels of primary roots by about 0 to 20%,while effects on nodal roots were slight. Generally, the ratioof cross-sectional areas between late and early metaxylem vesselsincreased from primary to seminal and nodal roots. Within thecultivar Tuxpefio this ratio was much reduced by M5. A few cultivarsmaintained the combined cross-sectional areas of metaxylem vesselsat M5 in some main root types, but only one cultivar could achievethis for the total of cross-sectional areas of metaxylem vessels,calculated over all root axes, by increasing the number of seminaland nodal roots. These anatomical traits seemed to be mostlyconstitutive with limited response to an actual environment,but they could be decisive for the suitability of a cultivarto an environment with frequent water shortages during seedlingestablishment. Key words: Metaxylem vessels, water stress, tropical maize     

Examination of morphological changes in the first formed protoxylem in Arabidopsis seedlings

We examined morphological changes in the first-formed protoxylem vessels in Arabidopsis seedlings. Between 2.5 and 8 days after imbibition, mean hypocotyl and root length increased 1.52 and 23.3 times, respectively. In the 2.5-day-old seedlings, two continuous protoxylem vessels were present in the hypocotyl-root axis. In the 8-day-old upper hypocotyls, six protoxylem vessels were observed, and in the lower hypocotyls, four protoxylem vessels and one or two metaxylem vessels were observed. In the 8-day-old roots, there were two protoxylem vessels and two or three metaxylem vessels. Two protoxylem vessels in the hypocotyls connected to two metaxylem vessels in the roots of 8-day-old seedlings. At the 0.3-mm part below the hypocotyl-root boundary, the mean intervals of neighboring annular secondary wall thickenings in protoxylem vessels in 8-day-old roots were 12.9% larger than those in 2.5-day-old roots. In more apical parts of 8-day-old roots, the mean intervals fluctuated between 1.71 and 2.29 μm. In 8-day-old seedlings, metaxylem vessels were formed between 0.4 mm above the hypocotyl-root boundary and 17 mm below the boundary. The intervals in these regions were not extended so much as protoxylem vessels were collapsed. The first-formed protoxylem vessels presumably retain their water-conductive function after metaxylem formation. Received: April 10, 2001 / Accepted: November 7, 2001     

Functional anatomy and xylem cavitation resistance of three species of monocotyledons grown on flooded substrates

There are few investigations that analyze the xylem functional anatomy of monocotyledons, as the methods have been developed for woody plants. This study describes the root, rhizome and aerial stem xylem anatomy and functional anatomy of Canna indica, Cyperus papyrus and Phragmites communis grown on flooded substrates; and it aims to evaluate the relationship between the xylem anatomy and its cavitation resistance. To calculate the indexes of vulnerability, mesomorphy, collapse and relative hydraulic conductivity in the three organs mentioned, the diameter, number of vessels per mm², thickness of the walls and the length of the tracheal elements were recorded. In addition, the xylem specific conductivity of the aerial stem was measured with the pipette method, and its resistance to cavitation was determined experimentally by the air injection technique. The protoxylem is xeromorphic, it has longer vessel elements, smaller diameters, thin walls and scalariform perforation plates, whereas the metaxylem is mesomorphic, with shorter vessel elements, larger diameters, thicker walls and simple perforation plates. Both present low collapse resistance but have a high relative hydraulic conductivity. P. communis recorded the highest cavitation resistance, and the number of vessels per mm² was related to xylem cavitation resistance in Canna indica and Cyperus papyrus. The experimental results of this investigation match partially the anatomical indexes and showed that the xylem of these species has a low specific conductivity and is more vulnerable to cavitation than that of other monocots.     

Subcellular localization of mineral deposits in the roots of wheat (Triticum aestivum L.)

Summary Mineral distribution in the roots of wheat (Triticum aestivum L. cv. Wheaton) was investigated using X-ray microanalysis of bulk frozen hydrated roots in SEM and of freeze substituted sections in TEM. Results obtained using the two methods agreed reasonably well. A total often elements were detected: Na, Mg, Si, P, S, Cl, K, Ca, Mn, and Fe. Of these Si, P, Ca, and Mn were incorporated into biomineralized structures. Silica was deposited in the endodermal walls in the older parts of the root. Silicon was also detected in the large central metaxylem lumina in the basal zone of the root, and in the smaller peripheral metaxylem and the immediately contiguous pericycle and outer parenchyma cells bridging the small metaxylem vessels to the endodermal layer. In the basal zone of the root some of the inner cortical cells contained intracellular electron opaque deposits. These were associated with the cell walls, had non-opaque inclusions and microanalysis revealed that they consisted of calcium, phosphorus and manganese.Abbreviations A apical zone of root - M midzone of root - B basal zone of root - SEM scanning electron microscope - TEM transmission electron microscope     

Effect of heavy metals and temperature on the oxalate oxidase activity and lignification of metaxylem vessels in barley roots

The effect of Cd on oxalate oxidase (OxO) activity and its localisation were analysed in barley root. In Cd-treated roots OxO activity was strongly induced in the region 2–4 mm behind the root tip and in the area toward the root base. In situ analyses showed that Cd-induced OxO activity was localised to the cell wall (CW) of early metaxylem vascular bundles and surrounding parenchyma cells and was accompanied by lignification of metaxylem vessels. OxO activation was also observed during treatment with other heavy metals (HMs), salt treatment and at elevated non-optimal temperature. In contrast to HM activation of OxO and lignification, high temperature and NaCl indeed activated OxO but did not induce lignification of metaxylem vessels. These results suggest that oxalate oxidase as an H₂O₂-generating enzyme is activated in response to several stresses, however the ectopic lignification of metaxylem vessels is activated specifically by HMs. This HM-induced premature root xylogenesis due to ectopic lignification of metaxylem vessels probably causes shortening of the root elongation zone and therefore a reduction in root growth.     

Xylem of rattans: vessel dimensions in climbing palms

We examined 11 species in four genera of rattans (Calamus, Daemonorops, Korthalsia, Plectocomia) growing in their native rainforest habitat in Singapore. Using aqueous safranin dye, we found that >95% of all vascular bundles at the base of a mature stem were functioning to transport water. We determined the frequency of vessel lengths in the long stems of these climbing palms by infiltration with dilute latex paint. Separate length distributions were made for metaxylem and protoxylem vessels; in both, there were many short and a few long vessels. The longest protoxylem vessels ranged from 7.5 to 62 cm in length, but one stem had an exceptional protoxylem vessel measuring 3.0 m. Maximum metaxylem vessel diameters were positively correlated to maximum vessel lengths in these species. The longest metaxylem vessel was found in K. rigida and was 3.96 m in length and was constructed from ～1200 vessel elements (cells). The widest vessel in that same stem was 532 μm in diameter. Long, wide vessels decrease resistance and increase water transport efficiency. In addition, we suggest that wide metaxylem vessels may have an important function in water storage.     

Ultrastructure of xylem parenchyma cells of barley roots in relation to ion transport to the xylem

Summary The structure of xylem parenchyma cells is examined in relation to transport of ions through the root. Measurement of uptake of ⁸⁶Rb⁺ and its transport through the root at different distances from the apex show that this is a general activity along the length of the root and not confined to a limited region. Thus transport through the root is not stopped by removal of that part of the root tip containing metaxylem vessels with living contents. The structure of xylem parenchyma appears to be suitable for involvement in ion transport from the stele to the xylem. At 1 cm behind the tip, where metaxylem vessels have no living contents but ion uptake and transport are going on at high rates, xylem parenchyma cells are rich in cytoplasm with extensive rough endoplasmic reticulum and well-developed mitochondria. Their cell walls contain numerous plasmodesmata, establishing the possibility of a symplastic pathway across the stele up to the vessels. The results are discussed in relation to regulation of ion transport to the xylem vessels in roots.Dedicated to Professor O. Stocker on the occasion of his 85th birthday.     

The Occurrence of Vascular Cavities and Specialized Parenchyma Cells in the Roots of Cool-season Legumes

The seeds of 20 legume species were grown in the greenhouse or in growth chambers at different temperatures. Under warm temperature conditions (above 15 °C), six species, pea (Pisum sativum), broadbean (Vicia faba), chickpea (Cicer arietinum), lentil (Lens culinaris), wild lupine (Lupinus latifolius), and soybean (Glycine max), formed cavities in the vascular cylinder of their primary roots, which in turn became filled by the ingrowth of specialized parenchyma cells (SP cells). When these species were grown at low temperature (below 15 °C), however, a “normal” vascular cylinder formed in the primary roots with late-maturing metaxylem vessel members differentiating in the center. These species were all cool-season legumes except soybean, a warm-season legume, which sometimes also formed cavities and SP cell ingrowths. The occurrence of cavities and SP cells therefore was restricted to the cool-season legumes (except soybean) when they were grown under warm temperature conditions. The position and size of cavities varied among these species. Pea and broadbean usually formed large, axially elongated cavities in the central vascular cylinder, or in the xylem poles. Others formed smaller cavities of various lengths.