Ultrastructure and Differentiation of Protein-storing Cells in Secondary phloem of Hevea brasiliensis Stem

The ultrastructure and differentiation of the protein-storingcells in secondary phloem of terminal branchlets of Hevea brasiliensisMull. Arg. were studied using electron microscopy. The cellsare parenchyma tissue in the axial system and characterizedby the presence of a large amount of proteinaceous fibrils inthe central vacuole. The fibrils of the protein-storing cellsare straight, about 9 nm in diameter and arranged in a directionroughly parallel to the longitudinal axis of the stem. At theearly differentiation stage of the protein-storing cells, amass of proteinaceous fibrils appears in the cytoplasm, thenis separated from the peripheral cytoplasm by the endomembranesystem derived from endoplasmic reticulum, resulting in theformation of the central vacuole with the fibrils inside asthe vacuolar content The peripheral cytoplasm may continue toproduce proteinaceous fibrils with which the fibrils of thecentral vacuole is supplemented after the protein-storing cellsare formed. Hevea brasiliensis, storage protein, proteinaceous fibrils, vacuole, secondary phloem     

Ultrastructure of compatible and incompatible interactions in phloem sieve elements during the stylet penetration by cotton aphids in melon

Resistance of the melon line TGR‐1551 to the aphid Aphis gossypii is based on preventing aphids from ingesting phloem sap. In electrical penetration graphs (EPGs), this resistance has been characterized with A. gossypii showing unusually long phloem salivation periods (waveform E1) mostly followed by pathway activities (waveform C) or if followed by phloem ingestion (waveform E2), ingestion was not sustained for more than 10 min. Stylectomy with aphids on susceptible and resistant plants was performed during EPG recording while the stylet tips were phloem inserted. This was followed by dissection of the penetrated leaf section, plant tissue fixation, resin embedding, and ultrathin sectioning for transmission electron microscopic observation in order to study the resistance mechanism in the TGR. The most obvious aspect appeared to be the coagulation of phloem proteins inside the stylet canals and the punctured sieve elements. Stylets of 5 aphids per genotype were amputated during sieve element (SE) salivation (E1) and SE ingestion (E2). Cross‐sections of stylet bundles in susceptible melon plants showed that the contents of the stylet canals were totally clear and also, no coagulated phloem proteins occurred in their punctured sieve elements. In contrast, electron‐dense coagulations were found in both locations in the resistant plants. Due to calcium binding, aphid saliva has been hypothesized to play an essential role in preventing/suppressing such coagulations that cause occlusion of sieves plate and in the food canal of the aphid's stylets. Doubts about this role of E1 salivation are discussed on the basis of our results.     

The evolution of minor vein phloem and phloem loading

Phylogenetic analysis provides a rational basis for comparative studies of phloem structure and phloem loading. Although several types of minor vein companion cell have been identified, and progress has been made in correlating structural features of these cells with loading mechanisms, little is known about the phylogenetic relationships of the different types. To add to the available data on companion cells, we analyzed the ultrastructure of minor veins in Euonymus fortunei and Celastrus orbiculatis (Celastraceae) leaves and determined that in these species they are specialized as intermediary cells. This cell type has been implicated in symplastic phloem loading. The data were added to published data sets on minor vein phloem characteristics, which were then mapped to a well-supported molecular tree. The analysis indicates that extensive plasmodesmatal continuity between minor vein phloem and surrounding cells is ancestral in the angiosperms. Reduction in plasmodesmatal frequency at this interface is a general evolutionary trend, punctuated by instances of the reverse. This is especially true in the case of intermediary cells that have many plasmodesmata, but other distinguishing characteristics as well, and have arisen independently at least four, and probably six, times in derived lineages. The character of highly reduced plasmodesmatal frequency in minor vein phloem, common in crop plants, has several points of origin in the tree. Thus, caution should be exercised in generalizing results on apoplastic phloem loading obtained from model species. Transfer cells have many independent points of origin, not always from lineages with reduced plasmodesmatal frequency.     

Hydraulic architecture of palms

Abstract

The water transport and storage system of palms is adapted to maintain the primary stem xylem functional over the life of the shoot, and in spite of severe drought. However, our structural information far exceeds our knowledge of vascular function, and these functional considerations bring more questions than answers. The tendency to generalize from limited data on a few species begs the question of how the hydraulic parameters discussed vary between palms with different growth forms and ecologies.     

The uniqueness of palms

Palms build tall trees entirely by primary growth in a way that limits their growth habit, but not their capacity for continued stem development. They achieve massive primary stature because of distinctive features of leaf development, stem vasculature and anatomical properties. They exhibit several record features of leaf and seed, and inflorescence size and leaves of great complexity. A marked ability to generate new roots allows them to be transplanted easily. As climbing plants they develop the longest unrooted stems in which there are, paradoxically, anomalous features of vascular construction compared with tree palms. It is here claimed that they are the world's longest lived trees because stem cells of several kinds remain active in differentiated tissues throughout the life of the palm. Absence of physiological dormancy may be related to this property, together with inability to withstand freezing temperatures that would cause irreversible cavitation of tracheary elements. This largely restricts them to the tropics, for which they are emblematic organisms. In these biological features palms are indeed unique organisms.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 5–14.     

Major trends of evolution in palms

From the diversity found among palms the following evolutionary trends are suggested:habit: from sympodial to monopodial;size: from moderate toward large and also toward small;stem: from unbranched to dichotomously branched, from little to much sclerenchyma, from short to elongate internodes;leaf: from an undivided eophyll to a palmate, costapalmate, pinnately ribbed or pinnate blade; from undivided and plicate to divided along the adaxial rib (“induplicate”) or along the abaxial rib (“reduplicate”); from pinnate to bipinnate or to pinnae onceor twicedivided longitudinally; from sheath split opposite the petiole to sheath tubular; from marcescent to deciduous; from central vascular bundles of the petiole with a single phloem strand to two phloem strands;inflorescence units: from moderately branched to spicate or less frequently to more diffusely branched, from one unit per leaf axil to more than one per axil, from among the leaves to below them or to above them in a compound terminal inflorescence, from pleonanthic to hapaxanthic;prophyll: from completely to incompletely encircling the peduncle, from incompletely to completely sheathing in bud;bracts: from conspicuous to small or absent at maturity, first peduncular bract from tubular and open at the apex to completely enclosing the inflorescence in bud, and then from ungrooved to deeply plicate;flower arrangement: from solitary, pedicellate, bracteolate flowers to a sympodial cincinnus of 2 or 3 or more, or to a short monopodial axis of 2–4 or more;bracteoles: from sheathing and prophyllate to completely closed or to incompletely developed or absent;flowers: from bisexual to unisexual, then associated with polygamy or monoecism to dioecism;perianth: from trimery to dimery or tetramery to decamery or to reduced and monochlamydeous;sepals: from distinct and imbricate to connate or separated;petals: from distinct and imbricate to valvate, or strongly imbricate, or connate; from small and ovate to large and variously shaped, or to small;stamens: from 6 to 3 or to more than 6 (to 950+);filaments: from relatively slender and distinct to broad and thick, and often connate or adnate to the perianth or both;staminodes: from stamenlike with abortive anthers only, to short teeth, or to a cupule at the base of the ovary, or to absent;pollen: from monosulcate to trichotomosulcate to dicolpate to monocolpate, diporate, or triporate;gynoecium: from apocarpous to syncarpous, from thin walls to thick, variously specialized walls;carpels or locules: from 3 to 2-1 or to 4–10;ovules: from moderate to small or to large, from anatropous to hemianatropous to campylotropous to orthotropous;pistillode: from only slightly modified from the gynoecium to vestigial or lacking or rarely to prominent;fruit: from fleshy to dry and fibrous;endocarp: from little differentiated or thin, to thick and hard, and sometimes with a pore or operculum over the embryo;seed: from moderate to small or to very large, from entire to dissected, bilobed, or perforate;endosperm: from homogeneous to invaginated or ruminate;germination: from remotetubular or -ligular to adjacent-ligular;chromosome complement: fromn = 18 ton = 17, 16, 15, 14, 13.     

Diversity of the superfamily of phloem lectins (phloem protein 2) in angiosperms

Phloem protein 2 (PP2) is one of the most abundant and enigmatic proteins in the phloem sap. Although thought to be associated with structural P-protein, PP2 is translocated in the assimilate stream where its lectin activity or RNA-binding properties can exert effects over long distances. Analyzing the diversity of these proteins in vascular plants led to the identification of PP2-like genes in species from 17 angiosperm and gymnosperm genera. This wide distribution of PP2 genes in the plant kingdom indicates that they are ancient and common in vascular plants. Their presence in cereals and gymnosperms, both of which lack structural P-protein, also supports a wider role for these proteins. Within this superfamily, PP2 proteins have considerable size polymorphism. This is attributable to variability in the length of the amino terminus that extends from a highly conserved domain. The conserved PP2 domain was identified in the proteins encoded by six genes from several cucurbits, celery (Apium graveolens), and Arabidopsis that are specifically expressed in the sieve element-companion cell complex. The acquisition of additional modular domains in the amino-terminal extensions of other PP2-like proteins could reflect divergence from its phloem function.     

Wound phloem in transition to bundle phloem in primary roots ofPisum sativum L.

Summary 48 hours after interrupting the root stele ofPisum, wound phloem initiated (proximally or distally to the wound) to reconnect the vascular stumps was found to contain some nucleate wound-sieve elements. At the elongating end of an incomplete wound-sieve tube these elements exhibit a sequence of ultrastructural changes as known from protophloem-sieve tubes. Elongation occurs by the addition of newly divided (wound-) sieve-element/companion-cell complexes. In order to dedifferentiate and assume a new specialization formerly quiescent stelar or cortical cells require at least one (mostly more) preliminary division. Companion cells are consequently obligatory sister cells to wound-sieve elements.By reconstruction using serial sections it could be shown that wound-sieve tubes elongate bidirectionally, starting in an early activated procambial cell of the stele. The elongation is directed by the existence of plasmodesmata, preferably when lying in primary pit fields, and by the plane of preceding divisions. Thus, the developing wound-sieve tube can deviate from the damaged bundle and radiate into the cortex as soon as the plane of the preceding divisions is favourable. In the opposite direction, elongating wound-sieve tubes run parallel to pre-existing phloem traces, thus broading their base at the bundle for the deviating part of the wound-sieve tube. Frequently an individual wound-sieve tube is supplemented at the bundle by a further wound-sieve tube which is partly running parallel to it. Both sieve tubes are interlinked with sieve plates by three-poled sieve elements.Ultrastructurally, the developmental changes of nucleate wound-sieve elements follow the known pattern. In spite of its contrasting origin and odd shape a mature wound-sieve element eventually has the same contents as regular sieve elements: sieve-element plastids, mitochondria, stacked ER and small amounts of P-protein within an electronlucent cytoplasm.     

New palms from Venezuela

Asterogyne yaracuyense andPrestoea humilis are described and illustrated, and their relationships discussed.     

Phytoferritin accumulations in leaves of diseased coconut palms

Summary A study by electron microscopy of coconut palm (Cocos nucifera L.) leaves from trees infected by the Cape St. Paul wilt (Kaincopé) disease of West Africa was carried out. Samples were obtained during the dry season (Dec.–Jan.) and fixed immediately upon removal from the trees in buffered glutaraldehyde. Further processing for electron microscopy was carried out within a week. No virus particles, mycoplasma-like organisms (MLO), fungi, or bacteria were detected in thin sections. Crystalline or paracrystalline accumulations of electron-opaque granules, approximately 5.5–6 nm in diameter, were observed in disintegrated chloroplasts of mesophyll cells. Based upon their morphological characteristics, formation of the slightly curved, fingerprint arrays or linear rows running parallel, and the visualization of electron-opaque cores in unstained preparations, the granules were identified as phytoferritin particles.This work was sponsored in part by National Science Foundation Grant GB-29280 and by a travel grant from the Food and Agriculture Organization of the United Nations.