STRUCTURE AND DEVELOPMENT OF SIEVE ELEMENTS IN THE LEAF OF ISOETES MURICATA

Leaf tissue of Isoetes muricata Dur. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. The very young sieve elements can be distinguished from contiguous parenchyma cells by their distinctive plastids and the presence of crystalline and fibrillar proteinaceous material in dilated cisternae of the rough ER. During differentiation, the portions of ER enclosing this proteinaceous substance become smooth surfaced and migrate to the cell wall. Along the way they apparently form multivesicular bodies which then fuse with the plasmalemma, discharging their contents to the outside. At maturity, the sieve element contains an elongate nucleus, which consists of dense chromatin material, and remnants of the nuclear envelope. In addition, the mature sieve element is lined by a plasmalemma and a parietal, anastomosing network of smooth ER. Both plastids and mitochondria are present. P-protein is lacking at all stages of development. Tonoplasts are. not discernible in mature sieve elements. The end walls of mature sieve elements contain either plasmodesmata or sieve pores or both, but only plasmodesmata occur in the lateral walls.     

SUBMICROSCOPIC STRUCTURE AND DEVELOPMENT OF THE CHLOROPLASTS OF PSILOTUM TRIQUETRUM

Sun , C. N. (Washington U., St. Louis, Mo.) Submicroscopic structure and development of the chloroplasts of Psilotum triquetrum. Amer. Jour. Bot. 48(4): 311–315. Illus. 1961.—Aerial stems and stem tips of Psilotum triquetrum were used for the study of the fine structure and development of chloroplasts. The chloroplasts of Psilotum are ellipsoidal, with a principal axis of approximately 13 μ and a short axis of approximately 3.6 μ. They are bounded by a well-defined outer membrane which consists of 2 layers. Within the laminar system of the stroma, the lamellated grana appear as sharply defined regions. The grana are about 1–1.6 μ in diameter. They are distributed more or less uniformly throughout the entire chloroplast with the exception of a very narrow peripheral zone. Relatively large, osmiophilic globules occur in groups in the stroma. The development of the Psilotum chloroplast may be summarized as follows: (1) in the undifferentiated proplastid, vesicles occur; (2) lamellated layers are formed by the fusion of vesicles; (3) the lamellae multiply by a process of thickening and splitting; (4) the grana are differentiated within a certain area by heterogeneous deposition of material and by further cleavage of the lamellae. Osmiophilic globules are present throughout the developmental stages, and increase in number and size with increase in age of the chloroplast.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF PSILOTUM NUDUM

Psilotum nudum (L.) Beauv. (Psilotopsida) has a simple, vascularized sporophyte with a dichotomously branching aerial axis. The number and lumen diameters of tracheids in the actinostele decrease in each subsequent branch, leading to an approximate halving of the measured hydraulic conductance (K_h) from segment to segment. To understand how the anatomy of P. nudum affects K_h, a biophysical model based on the Hagen-Poiseuille relation was developed that incorporated lumen diameter, tracheid taper, pit cavities, and pit membranes. Using a technique previously developed for ferns, pit membrane resistance was determined by measuring water flow before and after dissolving the pit membranes with cellulase. Measured K_h was in good agreement with K_h calculated with the model after excluding thick-walled late metaxylem tracheids that dye studies showed were nonconducting. Model simulations showed that the approximately 40% overlap observed for tracheids of P. nudum was in the range leading to greatest conductance and that K_h decreased to half for 20% overlap. The model also showed that the pit membranes account for an increasing percentage of total resistance to water flow as the lumen diameter increases. Thus, the removal of such primary wall material and the evolutionary origin of vessels would have substantially increased K_h.     

STRUCTURE AND DEVELOPMENT OF THE SIEVE ELEMENT IN THE STEM OF LYCOPODIUM LUCIDULUM

Stem tissue of Lycopodium lucidulum Michx. was fixed in glutaraldehyde and postfixed in osmium tetroxide for electron microscopy. Although their protoplasts contain similar components, immature sieve elements can be distinguished from parenchymatous elements of the phloem at an early stage by their thick walls and correspondingly high population of dictyosomes and dictyosome vesicles. Late in maturation the sieve-element walls undergo a reduction in thickness, apparently due to an “erosion” or hydrolysis of wall material. At maturity, the plasmalemma-lined sieve elements contain plastids with a system of much convoluted inner membranes, mitochondria, and remnants of nuclei. Although the endoplasmic reticulum (ER) in most mature sieve elements was vesiculate, in the better preserved ones the ER formed a tubular network closely appressed to the plasmalemma. The sieve elements lack refractive spherules and P-protein. The protoplasts of contiguous sieve elements are connected with one another by pores of variable diameter, aggregated in sieve areas. As there is no consistent difference between pore size in end and lateral walls these elements are considered as sieve cells.     

BIOMECHANICS OF PSILOTUM NUDUM AND SOME EARLY PALEOZOIC VASCULAR SPOROPHYTES

The elastic modulus E, flexural rigidity EI, static loading P, and volume fractions V of tissues of the aerial portions of Psilotum nudum are examined in terms of two Voigt models: 1) total elastic modulus E_T of a branch element is considered the additive property of the elastic modulus of tissues times their respective volume fraction, i.e., E_T = (EV), + (EV)₂ + … (EV)_n, and 2) E_T is considered the product of the elastic modulus of cell walls and the apoplastic volume fraction, i.e., E_T = E_cwV_a. The parameters EI and P, together with the length (l) of each branch element are combined into a dimensionless ratio, Pl²/EI (the load parameter L), to describe the relationship between static loading (Pl²) and the ability to sustain loading (EI). The load parameter of branch elements was found to decrease as the level of branching within aerial portions of this race of Psilotum was ascended. The magnitude of P also decreased acropetally. However, the decrease in L is primarily due to a disproportionate decrease in EI. The load parameter of the basalmost branch of aerial portions is smaller for larger aerial portions of this race of P. nudum. Thus, larger specimens are subtended by more rigid branch elements than smaller aerial portions. The Voigt models provide satisfactory approximations of the data from this race Psilotum and indicate that the principal load supporting tissues are the cortical sclerenchyma and other lignified tissues in older (lower branch elements). The mechanics of the three dimensional branching of P. nudum is extended to consider the aerial portions of some early Paleozoic vascular sporophytes (Algaophyton, Rhynia). Analyses indicate that biomechanical speculation about these fossils is potentially flawed by the ambiguity of anatomical data.     

ELECTROPHORETIC EVIDENCE FOR GENETIC DIPLOIDY IN PSILOTUM NUDUM

Psilotum nudum (2n = 104) has been considered an ancient polyploid, having resulted from repeated cycles of hybridization and allopolyploidy. However, electrophoretic analysis indicates that this species is genetically diploid despite its high chromosome number. Sixteen enzymes, encoded by 28 loci, revealed in P. nudum the number of isozymes typical of diploid seed plants. There is, therefore, no evidence of polyploid gene expression for the enzymes analyzed. These results for Psilotophyta are similar to those obtained for other lineages of homosporous pteridophytes, i.e., Arthrophyta and homosporous Microphyllophyta and Pteridophyta, all of which should be considered genetically diploid. Several hypotheses have been proposed to explain these results, most notably 1) cycles of allopolyploidy followed by massive gene silencing, and 2) initiation of these lineages with high chromosome numbers, possibly via chromosomal fission. Discrimination between these hypotheses awaits testing with molecular genetic techniques.     

THE FINE STRUCTURE OF SIEVE ELEMENTS OF NEREOCYSTIS LÜTKEANA

The sieve elements of Nereocystis from the base of phylloids contain numerous small vesicles, cytoplasm, ribosomes, and the usual organelles and membrane systems, including nuclei, plastids, mitochondria, dictyosomes, and endoplasmic reticulum. They have a thick secondary wall layer which is deposited along the longitudinal walls and at the sieve plate excluding the sieve pores. The sieve pores range in diameter from 100 to 400 nm and are lined by plasmalemma. The sieve elements from the hollow basal parts of the pneumatocyst show essentially the same features but have larger and fewer vesicles, relatively little cytoplasm, larger sieve pores, 400–900 nm in diameter, and may lack a nucleus. In old sieve elements there are large deposits of callose on the sieve plate and along the longitudinal wall; the vesicles seem to break down, and the protoplast appears necrotic. It is concluded that the trumpet hyphae and sieve tubes are basically the same type of cell, and that the trumpet-shape of the sieve elements is due to their passive stretching during extension growth of the organ in which they occur. There are minor but significant differences among the sieve elements from different regions of the thallus which may reflect possible levels of structural specialization of the sieve elements within the same plant.     

THE NACREOUS WALLS OF SIEVE ELEMENTS IN SEAGRASSES

The nacreous walls of sieve elements occur in seagrasses in all three genera of the family Zosteraceae and the genus Halodule of the family Cymodoceaceae but are absent from another eight seagrass genera belonging to the families Hydrocharitaceae, Cymodoceaceae, and Posidoniaceae. They occur in leaf blades, leaf sheaths, rhizomes, and erect stems but are not present in root tissues. The nacreous wall is uneven along the inner limits reflecting irregular thickness. The wall consists of hemicellulose or pectin and cellulose, but no protein, lignin, or lipid. Ultrastructurally, the wall contains parallel microfibrils or loose fibrils embedded in an amorphous matrix. Open pores occur in sieve plates and branching plasmodesmata are present in enlarged sieve areas. Mitochondria, endoplasmic reticulum, and plastids are also present in these sieve elements.     

STRUCTURE AND DEVELOPMENT OF THE SHOOT IN DOLICOTHELE

Boke , Norman H. (U. Oklahoma, Norman.) Structure and development of the shoot in Dolicothele. Amer. Jour. Bot. 48(4): 316–321. Illus. 1961.—A study of 2 species of Dolicothele reveals that although they have dimorphic areoles and a pattern of spine development similar to those of certain mammillarias, they share a significant number of ectomorphic and endomorphic characters with coryphanthas of the “vivipara group.” These include a tendency toward a cespitose habit; relatively large flowers; green fruits; pitted seeds; medullary vascular systems; forking of the main tubercle traces in the bases of the tubercles; lack of mucilage cells; thin-walled epidermis and hypodermis, both devoid of crystals; and large, druse-like crystalline aggregates in older parts of the pith and cortex. The evidence suggests that Coryphantha vivipara and closely allied species are the nearest extant relatives of Dolicothele. It would, therefore, seem inconsistent to return Dolicothele to Mammillaria unless an author's viewpoint were so conservative that he was willing also to return most, if not all, coryphanthas and escobarias to that genus.     

SLIME SUBSTANCE AND STRANDS IN SIEVE ELEMENTS

Studies of the secondary phloem of 6 species of woody dicotyledons revealed that slime is not normally dispersed throughout the vacuole of mature sieve elements, but occurs in the form of discrete strands that traverse the cell and run from cell to cell through the sieve-plate pores. As many as 5 fine strands, each measuring less than 0.5μ in diameter, were observed in a single pore. Less than 30% of pore area was occupied by strands. Thus, the pores are mostly open, and intervacuolar continuity exists between cells. These structural characteristics of pores offer strong support for the concept of mass flow.     

玉米叶片细脉原生韧皮部筛分子的分化和超微结构 : 原生质体的变化

玉米（ＺｅａｍａｙｓＬ．）叶片细脉原生韧皮部筛分子开始分化时,首先出现长的粗面内质网潴泡和增厚的细胞壁,随后在质体中出现其特征性的拟晶体内含物。随着分化的进行,长的粗面内质网潴泡转化为较短的形态,最后聚集成一些小的堆叠并失去其核糖体。细胞核发生退化,但常保持到成熟期的后期,此时的细胞核由双层核膜或某些部位仅由内核膜包被,内含电子致密的不定形染色质团块。随后双层核膜破裂而变得不连续。在细胞核开始退化时,核周腔局部膨大。有的膨大核周腔的外核膜破裂,并伴随邻近的部分细胞质解体。在核退化过程中,除内质网外,质体和线粒体的结构也发生变化,而核糖体、细胞质基质、液泡和高尔基体则解体消失。成熟原生韧皮部筛分子的原生质组分分布在细胞边缘,由质膜、线粒体、小的滑面内质网堆叠及具拟晶体的Ｐ型质体组成。随着邻近后生韧皮部筛分子分化的进行,成熟原生韧皮部筛分子的原生质组分逐渐退化,最后消失。     

紫云英花蜜腺的发育解剖学研究

紫云英花蜜除环绕在子房基部周围,属于花托蜜腺。成熟蜜腺的结构由分泌表皮和泌蜜组织构成。分泌表皮单层,细胞壁薄,其上有少量气孔存在,外切向壁角质层薄。泌蜜组织细胞5—6层。泌蜜组织基部具维管束、筛管或导管分子直达泌蜜组织基部。紫云英花蜜腺形成较晚,至露冠期基本成熟。PAS反应结果表明,紫云英花蜜腺是淀粉型蜜腺。蜜腺发育早期即在蜜腺细胞内充满众多的多糖颗粒。从结构和发育过程分析,紫云英花蜜腺的花蜜以分泌表皮层直接泌出为主,气孔泌出是辅助途径。