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.     

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.     

Sulla cariologia del Genere Carlina L.

Riassunto

L'A. ha studiato i cariogrammi di 6 specie italiane del genere Carlina L. (C. gummifera Less. var. Fontanesii (DC.) Fiori, C. acaulis L. var. alpina (Jacq.) Fiori, C. acanthifolia All., C. lanata L., C. vulgaris L., C. carymbosa L.).

Il corredo cromosomico è 2n = 20 per tutte le specie. Per la presenza di un gruppo di cromosomi della lunghezza superiore a 4 μ nelle specie a grossi capolini, si rende possibile distinguere auche cariologicamente nel genere Carlina L. due sezioni, corrispondenti alla possibile distinzione morfologica delle Carline in Macrocefale (C. gummifera Less. var. Fontanesii (DC) Fiori, C. acaulis L. var. alpina (Jacq.) Fiori, C. acanthifolia All., C. latana L.) e Microcefale (C. vulgaris L. e C. eorymbosa L.).     

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。     

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.     

Sporobolus Poibetii (R. et S.) Hitch. e Oenothera Sinuata L. Nella Selva di San Rossore

Summary

The author describes a Sporobolus Poiretii (R. et S.) Hitchc. and Oenothera sinuata L. stand in the «Selva» (forest) of S. Rossore (near Pisa).

Earlier reports of these two adventive and perhaps naturalized species are mentioned.

The author discussues the critical and complex sistematic definition, nomenclature and sinonimy of Sporobolus Poiretii in connection with the nearly related species Sporobolus indicus R. Br.

The author prospects at the end how both Sp. Poiretii and Oe. sinuata can be inserted in the «Nuova Flora Analitica» of Italy, by Adriano Fiori.     

How to become a tree without wood – biomechanical analysis of the stem of Carica papaya L

Carica papaya L. does not contain wood, according to the botanical definition of wood as lignified secondary xylem. Despite its parenchymatous secondary xylem, these plants are able to grow up to 10‐m high. This is surprising, as wooden structural elements are the ubiquitous strategy for supporting height growth in plants. Proposed possible alternative principles to explain the compensation for lack of wood in C. papaya are turgor pressure of the parenchyma, lignified phloem fibres in the bark, or a combination of the two. Interestingly, lignified tissue comprises only 5–8% of the entire stem mass. Furthermore, the phloem fibres do not form a compact tube enclosing the xylem, but instead form a mesh tubular structure. To investigate the mechanism of papaya's unusually high mechanical strength, a set of mechanical measurements were undertaken on whole stems and tissue sections of secondary phloem and xylem. The structural Young's modulus of mature stems reached 2.5 GPa. Since this is low compared to woody plants, the flexural rigidity of papaya stem construction may mainly be based on a higher second moment of inertia. Additionally, stem turgor pressure was determined indirectly by immersing specimens in sucrose solutions of different osmolalities, followed by mechanical tests; turgor pressure was between 0.82 and 1.25 MPa, indicating that turgor is essential for flexural rigidity of the entire stem.     

Evoluzione Annuale della Cerchia Legnosa in Individui Maschili E Femminili di Ginkyo Biloba L.

Abstract

Wood ring evolution in male and female plants of GINKYO BILOBA L. — A morphological examen of the wood rings in Ginkyo biloba L. has been performed on stem sections from specimens of the Botanical Institute of Florence, on wood samples from the collection of Adriano Fiori and on wood cores extracted from the stem of male and female plants growing in the Botanical Garden of Florence.

At the same time the development of the growth ring has been followed on wood samples tacken off the stems of two adult Ginkyo plants growing in the Botanical Garden of Florence.

All the material which has been studied shows the same characters, that is the frequency of uncomplete crescent-shaped wood rings, and consequently a large degree of eccentricity of the stems, depending not only upon the uncomplete wood rings, but also on the changing width of each ring.

Microscopical analysis has pointed out the presence of false rings, limited by discontinuous late wood, particularly in the female plant. This character has been observed only in a fossil wood of Ginkyo bonesii.

It has not been possible to assers how many rings or crescents are formed during one year. From dating records made on wood cores it seems that more than one ring could differentiate during one year, but the cambium shows a very irregular activity in Ginkyo. The factors which control the formation of false and uncomplete wood rings (wood crescents) are also unknown; their formation possibly depending upon environment conditions.

External factors such as water deficiency probably control cambial activity through auxin production. Larson (1964) has shown that draught causes a decrease in tracheids diameter, and the formation of false growth rings; thus the same causes could have been in action in our plants.

Noteworthy are also the data by Gungkel and Thimann (in Larson 1964) showing that auxin content increases downwards in the shoots of Ginkyo and that the apical bud has a very low auxin production, so that it seems possible that in Ginkyo the leaves supply something which the cortical tissue converts into auxin.

Lastly, the almost continuous growth activity of the cambium in Ginkyo must be pointed out, this causes a great difficulty in checking the beginning and the end of this process along the year. It is any way possible to establish the time of the highest cambial activity, which lies in June-July both in the female and in the male plants.     

Sul trasporto dei glucidi nelle piante superiori: I. — Aspetti biochimici

Abstract

CARBOHYDRATE TRANSLOCATION IN HIGHER PLANTS. I. - BIOCHEMICAL ASPECTS. — The concentration of soluble sugars and of hexose phosphates and the activity of several enzymes involved in hexose activation and polysaccaride synthesis have been investigated, separately, in the phloematic tissue and in the medullar parenchyma of Cucurbita Pepo internodes.

In the phloematic tissue (including sieve tubes, companion cells and phloematic parenchyma) the concentration of free hexoses appeared of about 50% lower, and that of glucose-6-P and of sucrose of about 100% higher then in the medullar parenchyma. Consistent amounts of raffinose were found only in the phloematic tissue. Paper chromatograms of the sieve tube exudate showed the presence of raffinose and sucrose in a ratio close to unity, and no appreciable amounts of free hexoses.

Determination of enzyme activity on preparations obtained from homogenates from the two types of tissue by repeated ammonium sulfate precipitation showed in the phloematic tissue a high activity of the enzymes hexokinase, UDP-kinase, UDPG-pyrophosphorylase and inorganic pyrophosphatase. The presence in the same tissue of galactosekinase, UDP-Gal-pyrophosphorylase and UDPG-epimerase was also ascertained.

On a protein basis, the activity of UDPG-pyrophosphorylase, inorganic pyrophosphatase and hexokinase appared about 3 times higher in the phloematic tissue than in the parenchyma; while this difference between the two tissues was not so marked for phosphofructokinase, and very small for other enzymes such as ATP-ase and phosphomono-esterase.

These results suggest that the very high activity, in the phloem cells neighbouring the sieve tubes, of the enzyme system catalyzing oligopolysaccaride synthesis could be an important component of the mechanism involved in the accumulation of oligopolysaccarides in the sieve tubes, and thus in sugar translocation. A scheme is proposed according to which the ATP and UTP energy would be utilized by the phloem cells to reach and to maintein a concentration of soluble sugars consistently higher than that prevailing in the contiguous tissues.     

药用植物商陆(Phytolacca acinosa Roxb.)根中异常次生结构的发育解剖学研究

本文报道了药用植物商陆根中异常次生结构的发生和发育过程。商陆根的初生结构和早期的次生结构都是正常的。但是,后来在维管柱的外围以离心的顺序先后产生5-7轮异常形成层.第一轮异常形成层起源于次生韧皮薄壁细胞和射线细胞。后一轮异常形成层在前一轮异常形成层向外产生的薄壁结合组织中发生。各轮异常形成层都以正常的活动方式产生同心环状排列的异常维管束以及它们之间丰富的薄壁结合组织,从而使根变成肉质状。薄壁结合组织细胞以及异常维管束内的薄壁组织细胞中贮藏有淀粉粒。     

Individual and combined effects of berberine and santonin on spore germination of some fungi

Berberine and santonin were isolated from rhizomes ofBerberis aristata and unexpanded flower buds ofArtemisia maritima, respectively. Efficacy of these two chemicals individually as well as of their mixtures, was tested against spore germination of some saprophytic and obligate fungi. While berberine individually was effective against most of the fungi,Helminthosporium spp. were least affected even at the highest dose (1500 ppm). Santonin was equally effective against several fungi. Mixture of both alkaloids found to be more effective than individual ones. Keeping the dose of berberine constant and santonin at two different concentrations (viz. 250 and 500 ppm) the spore germination ofHelminthosporium, oryzae was stimulated. Increasing concentration of santonin inhibited the spore germination of all other fungi tested,Colletotrichum capsici being affected only by 20 and 5% (at berberine concentration of 250 and 500 ppm, respectively). On the other hand, santonin being constant and berberine at different concentrations, the mixture was effective against all the fungi.     

Daphne thanguensis sp. nov. (Thymelaeaceae) from Sikkim Himalaya

Daphne thanguensis sp. nov. from north Sikkim of eastern Himalaya is described and illustrated. It is a narrow endemic related to D. tangutica Maxim. Daphne thanguensis grows in open alpine pastures and differs from D. tangutica by having leaves with revolute margin and a tuft of hairs at apex, ebracteate inflorescence and flowers, calyx lobes with a tuft of hairs at apex and annular, slightly undulate hypogynal disk. The new taxon is also close to D. retusa Hemsl. But can easily be differentiated by its sessile inflorescence, ebracteate flowers and tuft of hairs at leaf apices and calyx lobe apices. A conservation status of the new species in accordance with the IUCN red list categories and criteria is provided and discussed.     

Distichophyllum chenii B.C.Ho sp. nov. (Daltoniaceae) from Southeast China

Abstract

Despite the genus Distichophyllum having been revised recently for the Moss Flora of China, upon examination of a wide range of both Chinese and non-Chinese material a distinct new species was found among existing specimens collected from China. The new species, Distichophyllum chenii, is described based on specimens previously misidentified and reported as Distichophyllum cirratum var. cirratum. Published illustrations of D. cirratum var. cirratum based on a misidentified collection of D. collenchymatosum are highlighted to avoid further future taxonomic confusion.     

Formation of successive cambia and stem anatomy of Sesuvium sesuvioides (Aizoaceae)

Mature stems of Sesuvium sesuvioides (Fenzl) Verdc. were found to be composed of successive rings of xylem alternating with phloem. Repeated periclinal divisions in the parenchyma outside the primary phloem gave rise to conjunctive tissue and the lateral meristem that differentiate into the vascular cambium on its inner side. After the formation of the vascular cambium, the lateral meristem external to it became indistinct as long as the cambium was functional. As the cambium ceased to divide, the lateral meristem again became apparent prior to the initiation of the next cambial ring. The cambium was exclusively composed of fusiform cambial cells with no rays. In the young saplings, the number of cambial cylinders in the axis varied from the apex to the base, indicating formation of several rings within the year. In each successive ring of the lateral meristem, small segments differentiated into the vascular cambium and gave rise to vessels, axial parenchyma, fibres and fibriform vessels towards the inside, and secondary phloem on the outer side. In the old stems, non‐functional phloem of the innermost rings was replaced by a new set of sieve tube elements formed by periclinal divisions in the cambial segments associated with the non‐functional phloem. In some places the cambial segments completely differentiate into derivatives leaving no cambial cells between the xylem and phloem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 548–555.     

Ricerche Embriologiche su Senecio Vulgaris L. var. Thyrrenus Fiori

Abstract

Embryological researches on SENECIO VULGARIS L. var. THYRRENUS Fiori. — Male gametophyte, development of tapetal cells and female gametophyte have been studied in Senecio vulgaris L. var thyrrenus Fiori.

1) The development of male gametophyte is normal. Divisions of the microspore mother cells are of the simultaneous type. The division of the generative nucleus has never been observed till the pollen grain was in the anther.

2) The tapetal cells follow a very simple development. The nucleus of each cell divides only twice starting at the same time with the meiotic divisions of pollen mother cells but ending much earlier; subsequently, as usually happens with the Asteraceae, the ameboid involution of the tapetum begins. Endomitosis or any other process which leads to a polyploidy not due to nuclear fusion, has never been observed.

3) The female gametophyte is eight nucleate of the normal type (Polygonum). At maturity it shows only three antipodal cells whose nucleus undergoes at first, two or three divisions. Only later these new nuclei, always within the antipodal cell, may fuse in a polyvalent one.     

Cambial Activity and Distribution of Rubber in Guayule (Parthenium argentatum)

Vascular cambium in Guayule, a rubber producing Mexican shrubof Asteraceae family is non-storied. Cambial activity variesperiodically, and the vascular cambium and its immediate derivativesdo not contain rubber. However, as the xylem and phloem parenchymacells derived from the vascular cambium age, rubber depositionstarts from the cell periphery along the walls and later towardstheir cell lumen. Though the sieve tubes and companion cellsof phloem contain no rubber, all parenchyma cells of xylem andphloem, show the presence of rubber, though its amount varies.However, certain lignified xylem ray cells and lignified pithcells are devoid of rubber accumulation. Microfluorescence studiesshow that the epithelial, phloem ray parenchyma, cortical andpith cells, in descending order, have the highest to lowestrubber content. The size and number of rubber particles observedin the parenchyma cells are greatest during the period of cambialdormancy than in an active cambial period Cambium, guayule, rubber     

ANGIOSPERM WOOD FROM THE UPPER CRETACEOUS OF CENTRAL CALIFORNIA. III

Five species representing five genera are described. The specimens are part of a collection of woods from the Upper Cretaceous Panoche Formation of central California. Lardizabaloxylon cocculoides sp. n. resembles wood of those species of Cocculus of the Menispermaceae lacking included phloem. Carpinoxylon ostryopsoides sp. n. is similar to the wood of the Coryleae of the Betulaceae, particularly Ostryopsis. Riboidoxylon cretacea gen. et sp. n. is similar to the wood of Ilex of the Aquifoliaceae and also falls within the range of variation of the woods of the two closely related families, the Grossulariaceae and the Escalloniaceae. Mulleroxylon eupomalioides sp. n. falls well within the range of variation of the woods of the Magnoliaceae.     

On the glandular hairs of Cistus laurifolius L. (Cistaceae)

Abstract

Two types of glandular hairs have been found in Cistus laurifolius L. (Cistaceae). Both types present a great degree of morphological and histochemical variability, not only at population level but even within a single plant. This fact excludes that the hairs of C. laurifolius could be given a diagnostic value. The two hair types have a mixed hydrophilous and lipophilous secretion in which we have identified, especially in one type of them, cathecolic tannins. The powerful antimycotic activity of such tannins, makes the hairs particularly interesting for the possible ecological implications.     

The effect of different chiral morphs on visitation rates and fruit set in the orchid Spiranthes spiralis

Background: The arrangement of flowers on inflorescences is important for determining the movement of pollinators within the inflorescence and, consequently, the overall mating success and fruit set of a plant.

Aims: Spiranthes spiralis is an orchid that has a spiralled inflorescence. The species has two chiral forms that show opposite coiling directions (clockwise and anti-clockwise). We tested if this arrangement of inflorescence influences pollinator attraction and behaviour.

Methods: We surveyed two natural populations, analysed the reproductive compatibility of the two morphs and estimated pollination success in natural and experimental populations.

Results: We found that the two morphs were not isolated by pre- or post-mating barriers, occurred with a similar proportion in natural populations and showed similar levels of pollination success both in natural and artificial populations. However, we found a different pattern of pollination success along the inflorescences. In the two morphs, lower flowers experienced a higher pollination rate and this rate decreased along the inflorescence faster in anti-clockwise than in clockwise individuals.

Conclusions: This finding suggests that pollinators visit the flowers sequentially from the lower part of the inflorescences and leave the anti-clockwise individuals more rapidly than the clockwise ones. However, this pollinator behaviour is not detrimental for the pollination success of either of the two morphs.     

Sucrose partitioning between vascular bundles and storage parenchyma in the sugarcane stem: a potential role for the ShSUT1 sucrose transporter

A transporter with homology to the SUT/SUC family of plant sucrose transporters was isolated from a sugarcane (Saccharum hybrid) stem cDNA library. The gene, designated ShSUT1, encodes a protein of 517 amino acids, including 12 predicted membrane-spanning domains and a large central cytoplasmic loop. ShSUT1 was demonstrated to be a functional sucrose transporter by expression in yeast. The estimated K_m for sucrose of the ShSUT1 transporter was 2 mM at pH 5.5. ShSUT1 was expressed predominantly in mature leaves of sugarcane that were exporting sucrose and in stem internodes that were actively accumulating sucrose. Immunolocalization with a ShSUT1-specific antiserum identified the protein in cells at the periphery of the vascular bundles in the stem. These cells became lignified and suberized as stem development proceeded, forming a barrier to apoplasmic solute movement. However, the movement of the tracer dye, carboxyfluorescein from phloem to storage parenchyma cells suggested that symplasmic connections are present. ShSUT1 may have a role in partitioning of sucrose between the vascular tissue and sites of storage in the parenchyma cells of sugarcane stem internodes.