Sieve Pores Research Articles

A scalable, micropore, platelet rich plasma separation device

We have designed a novel, low energy platelet-rich-plasma (PRP) separator capable of producing 50 mL of PRP in 30 min, intended for military and emergency applications. Blood flows over a 3 mm length of sieve at high rates of shear. A plasma-platelet filtrate passes through the sieve's pores while erythrocytes remain. The filtrate is flowed over a second 3 mm length of smaller-pored sieve that withdraws plasma. Bulk blood volume is maintained by returning platelet-free plasma to the erythrocyte pool, enabling a nearly complete multi-pass platelet extraction. The total percentage of platelets extracted is:θ(T)=1-exp (-V(f)(T)Φ(P)/V) where V is the original plasma volume, V ( f )(T) is the total filtered volume, and ϕ ( P ) is platelet passage ratio (filtrate concentration/bulk average concentration) taken to be constant. Maximum θ(T) occurs at maximum V ( f )(T)× ϕ ( P ) Test microsieves, 3 mm long × 3 mm wide, were used. ϕ ( P ) values measured at various filtrate flow rates (20-100 uL/min) and utilizing various filter pore sizes (1.2-3.5 μm), was as high as 150 %. Maximum V ( f )(T)× ϕ ( P ) was achieved utilizing the 3.5 um filters at the highest flow rate, 100 uL/min. Erythrocyte leakages were always below 2,000/uL, far below the allowable limit stipulated by the American Association of Blood Banking. These data imply that a 13.7 cm(2) filter area is sufficient to achieve the target separation of 50 mL of platelet concentrate in 30 min. The filtration cartridge would consist of multiple microporous strips of 3 mm width arranged in parallel so that each element would see the conditions used in the prototype experiments presented here. Other microfiltration schemes suggest no method of scaling to practical levels.

Holocene chemical precipitates in the continental sabkha of Tayma (NW Saudi Arabia)

Studies on playas (known locally as inland sabkhat) are rare and lack a focus on their role as geoarchives for Holocene climate and environmental change. We present characteristic sediments and processes that illustrate the change from relatively humid to hyperarid conditions within the sedimentary record of the playa of Tayma (Saudi Arabia). The shift from the Holocene humid period (10–8 ka BP) to the hyperarid conditions of today left traces in this geoarchive. During the humid period, a perennial lake had formed in the playa depression. The occurrence of biota, especially ostracod valves and foraminifer shells, adapted to salinity fluctuations and physiochemical reactions of ostracods and foraminifers testify environmental changes. Phases of lake contraction and high salinities were responsible for the formation of ‘open water’ evaporites, such as aragonite needles, and ooids. Dilution occurred during rain events and flooding of the playa; it is characterised in the sedimentary infill by clastic layers that were deposited in graded sequences. The smooth surface, the phenomenon of microsomia and sieve pore variations of the ostracod valves provide evidence for changing salinities, which are testimony to short lived changes. These processes and further aridisation resulted in the formation of ‘capillary’ evaporites, which are dominated by sulphate minerals. The precipitates undergo fractionation and re-dissolution, which results in a typical stratification of salts with different solubilities.

Floating assembly of diatom Coscinodiscus sp. microshells

Diatoms have silica frustules with transparent and delicate micro/nano scale structures, two dimensional pore arrays, and large surface areas. Although, the diatom cells of Coscinodiscus sp. live underwater, we found that their valves can float on water and assemble together. Experiments show that the convex shape and the 40nm sieve pores of the valves allow them to float on water, and that the buoyancy and the micro-range attractive forces cause the valves to assemble together at the highest point of water. As measured by AFM calibrated glass needles fixed in manipulator, the buoyancy force on a single floating valve may reach up to 10μN in water. Turning the valves over, enlarging the sieve pores, reducing the surface tension of water, or vacuum pumping may cause the floating valves to sink. After the water has evaporated, the floating valves remained in their assembled state and formed a monolayer film. The bonded diatom monolayer may be valuable in studies on diatom based optical devices, biosensors, solar cells, and batteries, to better use the optical and adsorption properties of frustules. The floating assembly phenomenon can also be used as a self-assembly method for fabricating monolayer of circular plates.

Interaction between partitioning porous plate and rising bubbles in a trayed bubble column

In a trayed bubble column, the structure of the partitioning plate plays an important role on the bubble behavior. This study examined the effect of the opening ratio and pore size of the plate on the bubble break-up frequency and bubble size distribution. The sieve tray was used as the partitioning plate. The opening ratio was closely related to gas cap development. The stagnation of bubble flow and a gas cap were observed with an opening ratio less than 48.5%. The gas cap increased with decreasing opening ratio and increasing superficial gas velocity. The main effect of the sieve tray could be categorized into the additional drag force and bubble break-up depending on the sieve pore size. When the sieve pore size was smaller than the Sauter diameter of the bubble swarm, the movement of rising bubbles was interrupted by the drag force applied by the surrounding mesh lines. On the other hand, when the sieve pore size was larger than the Sauter diameter, the bubbles were affected by the additional bubble break-up. After the bubbles penetrated the sieve tray, the bubble size distribution shifted to a smaller one and the Sauter diameter decreased.

Modeling the hydrodynamics of Phloem sieve plates.

Sieve plates have an enormous impact on the efficiency of the phloem vascular system of plants, responsible for the distribution of photosynthetic products. These thin plates, which separate neighboring phloem cells, are perforated by a large number of tiny sieve pores and are believed to play a crucial role in protecting the phloem sap from intruding animals by blocking flow when the phloem cell is damaged. The resistance to the flow of viscous sap in the phloem vascular system is strongly affected by the presence of the sieve plates, but the hydrodynamics of the flow through them remains poorly understood. We propose a theoretical model for quantifying the effect of sieve plates on the phloem in the plant, thus unifying and improving previous work in the field. Numerical simulations of the flow in real and idealized phloem channels verify our model, and anatomical data from 19 plant species are investigated. We find that the sieve plate resistance is correlated to the cell lumen resistance, and that the sieve plate and the lumen contribute almost equally to the total hydraulic resistance of the phloem translocation pathway.

Callose deposition in the phloem plasmodesmata and inhibition of phloem transport in citrus leaves infected with “Candidatus Liberibacter asiaticus”

Huanglongbing (HLB) is a destructive disease of citrus trees caused by phloem-limited bacteria, Candidatus Liberibacter spp. One of the early microscopic manifestations of HLB is excessive starch accumulation in leaf chloroplasts. We hypothesize that the causative bacteria in the phloem may intervene photoassimilate export, causing the starch to over-accumulate. We examined citrus leaf phloem cells by microscopy methods to characterize plant responses to Liberibacter infection and the contribution of these responses to the pathogenicity of HLB. Plasmodesmata pore units (PPUs) connecting companion cells and sieve elements were stained with a callose-specific dye in the Liberibacter-infected leaf phloem cells; callose accumulated around PPUs before starch began to accumulate in the chloroplasts. When examined by transmission electron microscopy, PPUs with abnormally large callose deposits were more abundant in the Liberibacter-infected samples than in the uninfected samples. We demonstrated an impairment of symplastic dye movement into the vascular tissue and delayed photoassimilate export in the Liberibacter-infected leaves. Liberibacter infection was also linked to callose deposition in the sieve plates, which effectively reduced the sizes of sieve pores. Our results indicate that Liberibacter infection is accompanied by callose deposition in PPUs and sieve pores of the sieve tubes and suggest that the phloem plugging by callose inhibits phloem transport, contributing to the development of HLB symptoms.

Improvements on the Sieve Plate Structure of Movable Sieve Jig

Movable sieve jig has the characteristics of simple structure, low water consumption, large capacity, less infrastructure investment, and low production costs, which can replace manual rock picking and be used by many coal preparation plants. Due to the size and the shape of sieve pore, common sieve plate is easy to plug and has a short life with many materials leaking through the sieve, which leads to complex follow-up treatment process. Based on the analysis of coal preparation features, this paper proposes a structure of circular-wave sieve plate and conducts theoretical and experimental studies about that. The results show that the optimized circular-wave sieve plate can significantly improve the performance. Therefore, it is worth to be promoted in the industry.

Definitive Callose Deposition in Tapping Panel Dryness Affected Bark of Hevea brasiliensis

The structural and functional aspects of sieve elements in Tapping Panel Dryness (TPD) affected bark of Hevea brasiliensis, with special reference to definitive callose deposition, have been studied using histochemical methods. The effect of ethephon in definitive callose deposition in TPD affected bark also has been investigated, as the ethylene is reported to have a major role in TPD. The phloem of Hevea is characterized by sieve elements with compound sieve plates. The deposition of definitive callose on the sieve areas following TPD limits the translocation of sap through sieve elements. The amount of functional phloem was narrow and the accumulation of definitive callose was more in the sieve elements of TPD affected bark compared to that of unaffected bark of the same tree and of healthy trees. The affected area showed an increment in deposition of definitive callose in both vertical and horizontal extent of sieve areas; and there was a reduction in length of sieve plate, density of sieve pores, length of inter-sieve area wall, and number of sieve element rows in the inner bark. The lateral walls of non-functional sieve elements became thick and lignified. The anatomical changes in the sieve elements of TPD affected bark suggest that the translocation of phloem sap is regulated by deposition of definitive callose and less differentiation of sieve elements from vascular cambium. The external application of ethephon on bark enhanced the definitive callose deposition on sieve elements. The findings from the study reveal that ethylene plays a major role in the senescence of sieve elements.

The early Holocene humid period in NW Saudi Arabia – Sediments, microfossils and palaeo-hydrological modelling

An early to mid-Holocene humid period has been inferred from a variety of geo-bio-archives of the Sahara and the Arabian Peninsula including sabkhas, palaeo-lakes, sand dunes, wadis, speleothems or marine sediments. On the Arabian Peninsula, most of these records are located in the southern and southeastern part. Studies from the northern part are rare. This paper presents the first results from palaeo-environmental and hydrological investigations on the sabkha basin of the Tayma oasis, northwestern Saudi Arabia. Sedimentary characteristics, micro- and macrofauna, a digital elevation model based on DGPS measurements, and 14C-AMS data indicate the presence of a perennial lake with a minimum depth of 13 m, a stored water volume of 1.16 × 107 m3 and a surface of 18.45 km2 between 10,000–9000 cal BP. Foraminiferal test malformations and the shape of sieve pores on ostracod valves were used to detect trends in palaeo-salinity and ecological stress conditions. The reconstructed gradual contraction of the lake at least after 8500 cal BP reflects the long-term aridisation trend on the Arabian Peninsula after the early Holocene. Based on the hydrological water balance equation, quantitative data on minimum palaeo-rainfall during the early Holocene humid period were calculated. Input parameters for the equation are the minimum lake level, lake surface, and lake volume during the peak of the early Holocene humid period as well as palaeo-evapotranspiration, groundwater infiltration, and surface runoff. Accordingly, a perennial lake in the endorheic basin of the modern sabkha with a lake level at the same elevation as the uppermost shoreline deposit would have required a minimum annual precipitation of 150 ± 25 mm. This value amounts to c. 300% of recent precipitation rates.

Fine-crushing of Wood-cellulosic Material Modified for its Hydrolysis for Bioethanol Production

A prototype of a fine-crusher (grinder) of wood-cellulosic materials has been constructed with flail-knives and a rotary sieve rotating independently of each other. Pieces of bamboo in batches were fed into the fine-crusher, the crushed material was then treated with pressurized hot water (sub-critical water) to modify its structure and then hydrolyzed into glucose. With an increase in the diameter of the pores of the rotary sieve, the specific energy requirement for fine-crushing decreased remarkably from 500 to 250 kJ/kg, while the estimated glucose yield at 200°C decreased negligibly from 38% to 35%. The best rotational speed of the rotor and the diameter of the sieve pores were 1500 rpm and 4 mm, respectively.

Fine structure of phloematic trypanosomatid–coconut tree interaction

Phytomonas wilt or Hartrot is a fatal disease of palm (Arecaceae) species including Cocos nucifera (coconut) and is caused by a phloematic trypanosomatid, a promastigote parasite that inhabits phloem sieve elements of disease palms. In the present work, we described the morphology of the interaction between a phloematic trypanosomatid (Phytomonas staheli) and C. nucifera. Two varieties and one ecotype of the adult coconut palm from northeast and southeast Brazil were analyzed, totaling 34,000 plants. Coconut palm losses due to Hartrot varied according to the variety or ecotype and geographic area. Occurrence of Hartrot was insignificant in Rio de Janeiro state (southeast), but in Bahia state (northeast) losses were substantial when appropriate cultural practices were not applied. Symptomatic and healthy palm tissues were analyzed by light and electron microscopy. Laboratory diagnoses revealed the twisted promastigote form of phloematic trypanosomatids in the extracts of shoot apex, leaves, stems and inflorescence in diseased plants, but not in the healthy ones. No parasites were found in the roots. Although the general anatomy of healthy and diseased palms was similar, callose deposition in the sieve plates was revealed by histochemistry and immunocytochemistry in the diseased tissue. Plugging by the P-protein and plastid alterations was also observed. Our observations strongly suggest that parasite traffic between sieve elements took place, although their cell bodies were larger than the sieve pores. Phloematic trypanosomatid proliferation in the sieve tube elements might interrupt the transport of phloem or/and consume plant nutrients. In addition, an association between the percentage of sieve elements colonized by pathogen in palm tissues and disease severity was established.

Structure and Development of the Endophyte in the Parasitic Angiosperm Cuscuta japonica

The endophyte, that is, the haustorial part within the tissues of the host plant Impatiens balsamina, of the parasitic angiosperm Cuscuta japonica was studied with light and electron microscopy. The endophyte consisted mainly of vacuolated parenchymatous axial cells and elongate, superficial (epidermal) cells. Then the elongate, epidermal cells separated from each other and transformed into filamentous cells, called searching hyphae. The hyphae grew independently either intercellularly or intracellularly in the host parenchyma. The apical end of the hyphal cells was characterized by conspicuous, large nuclei with enlarged nucleoli and very dense cytoplasm with abundant organelles, suggesting that the hyphal cells penetrating host tissue were metabolically very active. Numerous osmiophilic particles and chloroplasts were noted in the hyphae. The osmiophilic particles were assumed to be associated with elongation of the growing hyphe. Plasmodemata connections between the searching hyphal cells of the parasite and the host parenchyma cells were not detected. Hyphal cells that reached the host xylem differentiated into water-conducting xylic hyphae by thickening of the secondary walls. A xylem bridge connecting the parasite and the host was confirmed from serial sections. Some hyphal cells that reached the host phloem differentiated into nutrient-conducting phloic hyphae. Phloic hyphae had a thin layer of peripheral cytoplasm with typical features of sieve-tube members in autotrophic angiosperms, i.e., parallel arrays of smooth endoplasmic reticulum, mitochondria, and plastids with starch granules. Interspecific open connections via the sieve pores of the host sieve elements and plasmodesmata of the parasite phloic hyphae were very rarely observed, indicating that the symplastic translocation of assimilate to the parasite from the host occurred.

Fine structure and X-ray microanalysis of silicified woods from a Tertiary basin Pohang, Korea by scanning electron microscopy

Anatomical descriptions are provided on silicified woods from a Tertiary basin Pohang, Korea by scanning electron microscopy and X-ray microanalysis. The silicified woods appeared to retain the original exterior morphology of the once grown trees, and exhibited various colors on the surface. As a component of the axial system in the secondary xylem, pores were oval to globose and measured approximately 200–300 μm in diameter in transverse planes. Seemingly bordered pits were also frequently observed in the tracheary elements. As a component of the axial system in the secondary phloem, sieve elements were found to have many sieve pores that were filled with numerous fine particles. In tangential planes, rays in the vascular cambium were approximately 500 μm long, and usually several cells wide (multiseriate). Meanwhile, several forms of microbial growth such as bacterial chains and hyphal growth of either fungi or actinomycetes were evident in the vessel lumens of unpolished silicified wood pieces. Some fracture surfaces were mainly characterized by the occurrence of polyhedral crystals, probably quartz, in the fissures. By X-ray microanalysis, iron was detected from the brown-colored regions, whereas calcium was also detected together with iron in the black-colored regions. Based on the rare occurrence of tracheids as the axial system in the secondary xylem, the silicified woods in this study can be intrinsically categorized into angiosperm groups in the region.

Evidence for functional heterogeneity of sieve element–companion cell complexes in minor vein phloem of Alonsoa meridionalis

Two modes of phloem loading have been proposed, apoplastic and symplastic, depending on the structure of sieve element-companion cell complexes (SE-CCCs) in minor vein phloem. Species are usually classified as either apoplastic or symplastic loaders although the cytology of SE-CCCs in minor veins of the majority of plants indicates that both mechanisms can be simultaneously involved in phloem loading. The functions of structurally different SE-CCCs in minor veins of the stachyose-translocating plant Alonsoa meridionalis were examined. A stachyose synthase gene, AmSTS1, was expressed in intermediary cells but not in the ordinary companion cell of the same vein. In contrast, sucrose transporter AmSUT1 protein was present in ordinary companion cells but not in the neighbouring intermediary cells. These data reveal the principles of phloem sap formation in A. meridionalis and, probably, in many other dicots. The two types of SE-CCCs within one and the same minor vein load different carbohydrates, using contrasting mechanisms for their delivery into the phloem. Lateral sieve pores in the minor vein phloem lead to mixing of the carbohydrates soon after loading. While symplastic and apoplastic pathways can function simultaneously during phloem loading, they are separated at the level of different SE-CCCs combined in phloem endings.

Response of Sweet Orange (Citrus sinensis) to ‘CandidatusLiberibacter asiaticus’ Infection: Microscopy and Microarray Analyses

Citrus greening or huanglongbing (HLB) is a devastating disease of citrus. HLB is associated with the phloem-limited fastidious prokaryotic alpha-proteobacterium 'Candidatus Liberibacter spp.' In this report, we used sweet orange (Citrus sinensis) leaf tissue infected with 'Ca. Liberibacter asiaticus' and compared this with healthy controls. Investigation of the host response was examined with citrus microarray hybridization based on 33,879 expressed sequence tag sequences from several citrus species and hybrids. The microarray analysis indicated that HLB infection significantly affected expression of 624 genes whose encoded proteins were categorized according to function. The categories included genes associated with sugar metabolism, plant defense, phytohormone, and cell wall metabolism, as well as 14 other gene categories. The anatomical analyses indicated that HLB bacterium infection caused phloem disruption, sucrose accumulation, and plugged sieve pores. The up-regulation of three key starch biosynthetic genes including ADP-glucose pyrophosphorylase, starch synthase, granule-bound starch synthase and starch debranching enzyme likely contributed to accumulation of starch in HLB-affected leaves. The HLB-associated phloem blockage resulted from the plugged sieve pores rather than the HLB bacterial aggregates since 'Ca. Liberibacter asiaticus' does not form aggregate in citrus. The up-regulation of pp2 gene is related to callose deposition to plug the sieve pores in HLB-affected plants.

Plant- and stimulus-specific variations in remote-controlled sieve-tube occlusion

Phloem injury triggers local sieve-plate occlusion including callose-mediated constriction and protein plugging of sieve pores. In intact plants, reversible sieve-plate occlusion is induced by electric potential waves (EPWs)—accompanied by Ca2+-influx—as result of distant burning. Here, we present additional results which pertain to (a) the variability of EPW-profiles in relation to forisome conformation in intact Vicia faba plants and (b) the differential occlusion reactions to burning and cutting in various plant species. A correlation between stimulus perception and mode of phloem loading is discussed.Addendum to: Furch ACU, Hafke JB, Schulz A, van Bel AJE. Ca2+-mediated remote control of reversible sieve-tube occlusion in Vicia faba. J Exp Bot 2007; 58:2827-38.

A Plasma Membrane-Anchored Fluorescent Protein Fusion Illuminates Sieve Element Plasma Membranes in Arabidopsis and Tobacco

Rapid acquisition of quantitative anatomical data from the sieve tubes of angiosperm phloem has been confounded by their small size, their distance from organ surfaces, and the time-consuming nature of traditional methods, such as transmission electron microscopy. To improve access to these cells, for which good anatomical data are critical, a monomeric yellow fluorescent protein (mCitrine) was N-terminally fused to a small (approximately 6 kD) membrane protein (AtRCI2A) and stably expressed in Arabidopsis thaliana (Columbia-0 ecotype) and Nicotiana tabacum ('Samsun') under the control of a companion cell-specific promoter (AtSUC2p). The construct, called by its abbreviation SUmCR, yielded stable sieve element (SE) plasma membrane fluorescence labeling, even after plastic (methacrylate) embedding. In conjunction with wide-field fluorescence measurements of sieve pore number and position using aniline blue-stained callose, mCitrine-labeled material was used to calculate rough estimates of sieve tube-specific conductivity for both species. The SUmCR construct also revealed a hitherto unknown expression domain of the AtSUC2 Suc-H(+) symporter in the epidermis of the cell division zone of developing root tips. The success of this construct in targeting plasma membrane-anchored fluorescent proteins to SEs could be attributable to the small size of AtRCI2A or to the presence of other signals innate to AtRCI2A that permit the protein to be trafficked to SEs. The construct provides a hitherto unique entrée into companion cell-to-SE protein targeting, as well as a new tool for studying whole-plant phloem anatomy and architecture.

Anatomical descriptions of silicified woods from Madagascar and Indonesia by scanning electron microscopy

Fine structure and tissue substitution by minerals were investigated in silicified woods from Madagascar and Indonesia by scanning electron microscopy and X-ray microanalysis. The silicified woods maintained the exterior morphology of once grown trees and showed typical inner structures of conifers. Radial planes of the silicified wood from Madagascar revealed tracheids as a major component of the axial system in the secondary xylem. Tracheids were mainly characterized by numerous bordered pits where a thickening in the middle (torus) was surrounded with the membrane (margo). The torus appeared to contrast with the fibrillar network of the margo. As a component of the axial system in the secondary phloem, sieve elements were found to have many sieve pores that were filled with seemingly crystalline materials. To correlate the colors of the silicified wood from Indonesia with elemental composition, energy-dispersive X-ray spectrometry was employed in this study. Silicon was present as a basic component of the silicified wood. Calcium and iron were detected from red-colored regions, whereas magnesium was found in blue-colored regions. These results suggest that tissues of silicified woods had been substituted by minerals over the past period, while retaining the inherent morphology of the tree species. Scanning electron microscopy and X-ray microanalysis could be applied to unravel structural details and composition of plant fossils in palaeobotany.

Ca2+-mediated remote control of reversible sieve tube occlusion in Vicia faba

According to an established concept, injury of the phloem triggers local sieve plate occlusion including callose-mediated constriction and, possibly, protein plugging of the sieve pores. Sieve plate occlusion can also be achieved by distant stimuli, depends on the passage of electropotential waves (EPWs), and is reversible in intact plants. The time-course of the wound response was studied in sieve elements of main veins of intact Vicia faba plants using confocal and multiphoton microscopy. Only 15-45 s after burning a leaf tip, forisomes (giant protein bodies specific for legume sieve tubes) suddenly dispersed, as observed at 3-4 cm from the stimulus site. The dispersion was reversible; the forisomes had fully re-contracted 7-15 min after burning. Meanwhile, callose appeared at the sieve pores in response to the heat shock. Callose production reached a maximum after approximately 20 min and was also reversible; callose degraded over the subsequent 1-2 h. The heat induction of both modes of occlusion coincided with the passage of an EPW visualized by electrophysiology or the potential-sensitive dye RH-414. In contrast to burning, cutting of the leaf tip induced neither an EPW nor callose deposition. The data are consistent with a remote-controlled occlusion of sieve plates depending on the longitudinal propagation of an EPW releasing Ca(2+) into the sieve element lumen. It is hypothesized that forisome plugs and callose constriction are removed once the cytosolic calcium level has returned to the initial level in those sieve tubes.

Secondary phloem anatomy of Cycadeoidea (Bennettitales)

Secondary phloem anatomy of several species of Cycadeoidea is described from trunks in the Wieland Collection, Peabody Museum of Natural History. The trunks were collected from the Lakota Formation, Lower Cretaceous, Black Hills of South Dakota. Secondary phloem is extensively developed and consists of alternating, tangential bands of fibers and sieve elements, with rare phloem parenchyma. Uniseriate rays, 2-22 cells high, occur between every one to three files of the axial system. Fibers are long, more than 1200 μm, approximately 26.6-34.2 μm in diameter, and have slit-like apertures on the lateral walls. Sieve elements range from 16-25 μm in diameter and are up to 500 μm long. Elliptical sieve areas appear on both end and radial walls and measure 10 μm across; minute spots, which may represent sieve pores, are present within the sieve areas. Secondary phloem of North American Cycadeoidea is similar in organization (alternating tangential bands) and cell types (sieve cells, fibers, axial parenchyma) to that known in other extant and fossil cycadophytes and some seed ferns. The unusual pattern of cell types and thickness of secondary phloem is discussed in the context of plant habit, phloem efficiency, and potential phylogenetic importance.