The sieve tube wall and its relation to translocation

Summary The sieve tube wall possesses a broad inner layer often with pronounced radial striations. The plasmalemma of the sieve tube appears to penetrate this wall in the form of a brush border of irregular microvilli, greatly increasing its surface area. It is suggested that this is the site of active transport of potassium, which circulates electroosmotically through the sieve plate pores and back through the thick wall. The function of the companion cells is the care and maintenance of the active brush border sites; in conjunction with their activity in supplying high-energy intermediates movement in the column acts regeneratively and fully polarises the plates. Many of the lamellar stacks and curvilinear membrane aggregates hitherto regarded as endoplasmic reticulum are, it is suggested, plasmalemma displaced from the wall. These findings have important consequences for the electroosmotic theory.     

Evolutionary epistemology as an overlapping,interlevel theory

I examine the branch of evolutionary epistemology which tries to account for the character of cognitive mechanisms in animals and humans by extending the biological theory of evolution to the neurophysiological substrates of cognition. Like Plotkin, I construe this branch as a struggling science, and attempt to characterize the sort of theory one might expect to find this truly interdisciplinary endeavor, an endeavor which encompasses not only evolutionary biology, cognitive psychology, and developmental neuroscience, but also and especially, the computational modeling of artificial life programming; I suggest that extending Schaffner's notion of interlevel theories to include both horizontal and vertical levels of abstraction best fits the theories currently being developed in cognitive science. Finally, I support this claim with examples drawn from computational modeling data using the genetic algorithm.     

Proliferating sieve elements present in bud phloem anastomoses connect sieve tubes of axillary bud traces to stelar vascular bundles in the aquatic monocotyledonPotamogeton natans L. (Potamogetonaceae)

Summary The stem ofPotamogeton natans is characterized by a central stelar vascular system with reduced xylem and abundant phloem. Wide sieve tubes composed of short sieve-tube members joined by simple sieve plates and associated with companion cells establish an effective conduit for assimilates. At each node the phloem forms a network of parallel sieve elements connecting the stem phloem to leaf and bud traces. InP. natans an axillary bud rarely develops into a side branch, its procambial vascular bundles are each connected to the nodal complex via separate anastomoses. Their most unusual components are the anastomosai sieve elements (ANSE), characterized by thin cell walls pitted all over by tiny callose-lined pores resembling plasmodesmata, which can be detected as bright areas by fluorescence microscopy after staining with aniline blue. Several layers of ANSE make up the centre of an anastomosis and link to both the nodal and bud stelar sieve tubes via mediating (MSE) and connecting sieve elements (CSE). The ultrastructural differentiation of ANSE, MSE, and CSE corresponds to that of normal sieve elements, i.e., in the mature stage they are enucleate, evacuolate, and have lost most of their cytoplasm. Their plastids are of form-P2c, containing many cuneate protein crystals, typical of monocotyledonous sieve elements. Quantitative aspects of the pore areas are discussed in relation to the functional significance of bud anastomoses.Abbreviations ANSE anastomosai sieve elements - CSE connecting sieve elements - FM fluorescence microscopy - LM light microscopy - MSE mediating sieve elements - TEM transmission electron microscopy Dedicated to Professor Dr. Rainer Kollmann on the occasion of his retirement     

Histological localization of citrus infectious variegation virus (CVV) inPhaseolus vulgaris

Summary Ultrathin sections ofPhaseolus vulgaris leaves were studied in the electron microscope. The leaves were taken from plants, both healthy and experimentally infected with CVV. The sieve tubes and companion cells of all samples contained a slime-like substance, more or less organized into compact systems. In the mature sieve elements of virus-infected plants systems of parallel membranes were seen along which spherical particles, of about 30 m diameter, were aligned in simple rows.Pubbl No. 117 of Gruppo di Ricerca per le Virosi, del C. N. R.     

Metabolism of free and membrane-bound ribosomes during aging of Jerusalem artichoke tuber slices

Summary A biochemical and cytochemical study has been made of the distribution of -glycerophosphatase (EC 3.1.3.2) activity in mature and differentiating phloem cells of Nicotiana tabacum L. and the pH dependence and kinetics of -glycerophosphate hydrolysis of homogenates of fresh leaf midveins and midveins fixed in formaldehyde-gluteraldehyde. -glycerophosphatase showed two peaks of activity at pH 5.5 and 6.2. Enzyme saturation kinetics were exhibited by both fresh and fixed tissue homogenates. At a substrate concentration of 2 mM, 65% of the enzyme activity survived fixation. Specimens for cytochemical localization were incubated with 2 mM -glycerophosphate at pH 5.5 and 6.2. Specimens showed consistent patterns of reaction product deposition. Little or no reaction product was deposited in controls incubated without substrate or with substrate plus 0.01 M fluoride. -glycerophosphatase activity in the phloem and xylem is considerably higher than in surrounding tissue. Dense localization of reaction product was demonstrated on the vacuolar membranes, the inner membranes of mitochondria, and the dictysomes of phloem parenchyma and companion cells. The plasma membrane and endoplasmic reticulum cisternae of these cells were usually free of reaction products. Enzyme activity in mature sieve elements was associated with the parietal and stacked systems of endoplasmic reticulum and with the P-protein. There was inconsistency of staining of P-protein in mature sieve elements although the association of reaction products with the P-protein appeared to show a correlation with maturity and dispersal. The P-protein bodies of differentiating sieve elements showed no reaction product deposition. The distribution of -glycerophosphatase activity has been compared with that previously recorded for ATPase activity in the phloem of Nicotiana tabacum.     

Protocol-dependence of equivalent circuit parameters of toad urinary bladder

Summary Determinations of current-voltage relationships are widely employed in the characterization of epithelial sodium transport. In order to determine the protocol dependence of transport parameters in the toad urinary bladder, studies were carried out in the presence and absence of amiloride, an inhibitor of active sodium transport. With symmetric positive and negative perturbations of the transepithelial electrical potential difference (0±100 mV) for 30 sec, the amiloride-sensitive current-voltage (i ^a -) relationship was near linear over the range –75+100 mV, indicating constancy of the conductance ^a and the apparent electromotive force E _Na, lumped parameters of the standard electrical equivalent circuit model of the active transport system. With a reverse protocol (±1000 mV) or 15 min perturbations thei ^a - relationships were highly nonlinear. Nonlinearity reflected voltage dependence of parameters: perturbations that increased active transport decreased E _Na and increased ^a, as evaluated from 10 sec perturbations of ; slowing of active transport produced the converse changes. These effects are usefully analyzed in both quasi-steady states and true steady states by means of a detailed equivalent circuit incorporating the significant ionic currents across each plasma membrane. Precise understanding of the significance of ^a and E _Na will require characterization of the partial ionic conductances on perturbation of .     

Electron microscopic and cytochemical study on the role of Golgi elements and plasma membrane of enterocytes in the intestinal lipid transport

Summary Intestinal lipid absorption and transport were investigated in albino rats. The observations point towards the existence of a continuity between plasma membrane invaginations and elements of the Golgi complex on its mature face. They also suggest a segregation of lipid droplets by paired Golgi membranes and plasma membrane invaginations. The following way for lipid transport is deduced: lipid droplets moving inside the smooth endoplasmic reticulum accumulate progressively and are condensed in Golgi cisternae of the forming face. Their limiting membrane ruptures and liberated lipid droplets are segregated by paired Golgi membranes of the mature face or by plasma membrane invaginations. Subsequently the inner of the two segregating membranes disappears while the lipid droplet is moved towards the intercellular space inside a canal communicating with this space. The suggestion is made that the Golgi apparatus is of double origin: one component representing a terminal plication of the endoplasmic reticulum; the second one—a terminal plication of the plasma membrane invagination. This concept explains the ultrastructural and histochemical differences between Golgi membranes of the forming and mature faces of the complex.     

Über den Feinbau „Gitterartig” Aufgebauter Plasmaeinschlüsse in den Siebelementen von Dioscorea Reticulata

Zusammenfassung Ein hochgeordnetes System tubulärer Strukturen wurde als bisher unbekannter Plasmaeinschluß in den Siebelementen von Dioscorea reticulata beschrieben. 250 Å weite, kontrastreiche Tubuli ordnen sich in hexagonal dichtester Packung zu 0,2 bis 1,3 großen Körpern an. Die einzelnen Tubuli sind helixähnlich gewunden und geben dem Plasmaeinschluß eine den Heitz-Leyon-Kristallen der Plastiden von Chlorophytum ähnliche Gestalt. Herkunft und Bedeutung dieser Gebilde sind noch weitgehend unbekannt.

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On the fine structure of a lattice-like body in the sieve elements of Dioscorea reticulata

Summary In the parietal protoplast of differentiated sieve elements of Dioscorea reticulata an up to now unknown lattice-like body has been described in detail. Many 250 Å wide tubules combine to form very regularly composed bodies, which are about 0.2 to 1.3 large. Each tubule is twined like a helix and is spaced equidistantly from the adjoining ones, the interval being 450 to 500 Å. On the whole the lattice-like bodies of Dioscorea largely resemble the so-called Heitz-Leyon-crystals of young plastids.

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Mit 2 Textabbildungen     

Potassium permanganate fixative and the electron microscopy of sieve tube contents

Summary Sieve tube slime is probably fibrillar as suggested by other workers. The granular material seen in electron micrographs of sieve tubes fixed with potassium permanganate is mostly a precipitate produced by the reaction of the permanganate with the sieve tube contents, particularly with sucrose, and with reagents used in the fixing, dehydrating and embedding process. Potassium permanganate is not therefore a good fixative for the electron microscopy of sieve tube contents. Neither is it suitable for the study of fine structure in vacuoles or vesicles containing precipitable solutes in other kinds of cells nor for examining the fine structure of their ground cytoplasm.     

Notes on the Mechanism of ATP Synthesis

The most commonly quoted mechanism of the coupling between the electrochemical proton gradient and the formation of ATP from ADP and P_i assumes that all states of the F₁ portion of the ATP synthase have subunits in tight, loose, and open conformations. Models based on this assumption are inconsistent with some of the available experimental evidence. A mechanism that includes an additional subunit conformation, closed, observed in the rat liver structure overcomes these difficulties.     

Methodological problems in evolutionary biology VIII. Biology and culture

Biology cannot accommodate all aspects of culture. Aspects of culture that a biological approach can take into account can be covered by the biological categories of phenotype and environment. There is no need to treat culture as a separate category. Attempts to elaborate biological explanations of cultural variation will meet with success only if biologists expand theories of development, and integrate them in evolutionary biology. The alternative — elaborating the idea of so-called cultural inheritance — makes little sense from a biological point of view.     

The distribution of P-Protein in mature sieve elements of celery

Summary The extent of blocking of sieve-plate pores caused by release of cell turgor was investigated by fixing and processing for electron microscopy a long length of celery (Apium graveolens L.) phloem. Differences in distribution of P-protein within the pores were observed between those cells near the two cut ends, and the central cells.To assess the effect of chemical fixation on the distribution of P-protein, strands of celery phloem (fixed or unfixed, and not treated with cryoprotectants) were frozen in Freon 12 and then freeze-substituted. In sieve elements from unfixed tissue there were a greater number of sieve plates displaying partially open pores.Direct freezing of unprotected phloem tissue in Freon 12 resulted in the formation of ice crystals within the lumen of the sieve elements. Freezing of tissue at rates fast enough to avoid the formation of damaging ice crystals resulted in sieve-plate pores having an unoccluded central channel with a peripheral lining of P-protein. In the lumen of the sieve elements the P-protein filaments occurred as discrete bundles ca. 0.5 m in diameter, and as a parietal layer varying in thickness from 0.1 to 0.5 m.     

Feet,bridges, and pillars in triad junctions of mammalian skeletal muscle: Their possible relationship to calcium buffers in terminal cisternae and T-tubules and to excitation-contraction coupling

Summary The structure of the triad junction was examined in thin sections of mammalian fast-twitch skeletal muscle. The aims of the experiments were twofold: first, to examine relationships between the contents of the junctional gap and the terminal cisternae that could be significant in excitation-contraction coupling and, second, to look for structures in the transverse tubules that could support a calcium buffer system. Procedures known to stabilize cytoskeletal elements were used in an attempt to retain the original structure. Feet, pillars and bridges were often seen side by side in the same junction. In one such junction, the average center-to-center spacing between four bridges was 30.9±1.7 nm and between five foot-like structures was 29.2±1.4 nm. The subunit structure of the feet could be seen in many sections. The lumen of the terminal cisternae was filled with a tetragonal network of calsequestrin which formed parallel strands near the junctional membrane, in register with the feet. The strands overlay the area occupied by rods seen in freeze-fracture replicas of terminal cisterna membrane. The contents of the transverse tubules were aggregated into bands, or tethers, which extended across the short axis of the tubule at regular intervals of about 30 nm. The tethers consisted of flattened discs, stacked across the long axis of the tubule, aligned with the junctional feet. Lanthanum staining of the tethers indicated cationic binding sites that could buffer luminal calcium ion concentration in the vicinity of the voltage sensor for contraction. It is suggested (i) that the control of calcium concentration near the voltage sensor is necessary for normal activation, (ii) that feet, pillars and bridges are different images of a spanning structure, and (iii) that the regular alignment of tethers, feet and calsequestrin is functionally significant in excitation-contraction coupling.     

Endoplasmic reticulum and crystalline fibrils in the root protophloem of Nymphoides peltata

Endoplasmic reticulum in the root protophloem of Nymphoides peltata (S.G. Gmel.) O. Kuntze changes form as sieve elements differentiate. In immature sieve elements the individual endoplasmic reticulum (ER) cisternae form large irregular aggregates in the cytoplasm. In older immature sieve elements the ER aggregates are more ordered and membranes in them are convoluted. Although convoluted ER predominates in immature sieve elements the ER of the mature sieve elements consists mainly of flattened stacks of ER cisternae. Some of these stacks of ER may be derived from the existing convoluted ER. Crystalline fibrils first appear in the cytoplasm of the sieve element when the ER starts to aggregate. The crystalline fibrils move to the parietal layer of the sieve element along with the aggregates of ER. A possible ontogenetic relationship between ER and crystalline fibrils is discussed.Abbreviation ER endoplasmic reticulum     

Ultrastructural indications for coexistence of symplastic and apoplastic phloem loading in Commelina benghalensis leaves

The ultrastructural ontogeny of Commelina benghalensis minor-vein elements was followed. The mature minor vein has a restricted number of elements: a sheath of six to eight mestome cells encloses one xylem vessel, three to five vascular parenchyma cells, a companion cell, a thin-walled protophloem sieve-tube member and a thick-walled metaphloem sieve-tube member. The protophloem sieve-tube member (diameter 4–5 m; wall thickness 0.12 m) and the companion cell originated from a common mother cell. The metaphloem sieve-tube member (diameter 3 m; wall thickness 0.2 m) developed from the same precursor cell as the phloem parenchyma cells. Counting the plasmodesmatal frequencies demonstrated a symplastic continuum from mesophyll to the minor-vein phloem. The metaphloem sievetube member and the phloem parenchyma cells are the termini of this symplast. The protophloem sieve-tube member and companion cell constitute an insulated symplastic domain. The symplastic route, mesophyll to metaphloem sieve tube, appears to offer a path for symplastic loading; the protophloem sieve tube may be capable of accumulation from the apoplast. A similar two-way system of loading may exist in a number of plant families. Plasmodesmograms (a novel way to depict cell elements, plasmodesmatal frequencies and vein architecture) of some other species also displayed the anatomical requirements for two routes from mesophyll to sieve tube and indicate the potential coexistence of symplastic and apoplastic loading.     

Quantitative relationship between active sodium transport,expansion of endoplasmic reticulum and specialized vacuoles (“scalloped sacs”) in the outermost living cell layer of the frog skin epithelium (Rana temporaria)

Summary When an isolated frog skin (Rana temporaria) is exposed to a hydrostatic pressure difference between inside and outside bathing solutions (inside pressure higher than outside) of 20–50 cm of H₂O and if under these conditions the skin is short-circuited electrically, small vacuoles appear light-microscopically in the outermost living cell layer in the epithelium. The number of such vacuoles shows a linear dependency on the rate of active sodium transport as measured by the short-circuit current. Electron-microscopically, the vacuoles are interpreted as previously undescribed organelles, the scalloped sacs which are about 0.5 in diameter, with a wrinkled surface and bounded by a unit membrane. This organelle is in intimate contact with sacs and tubules of smooth endoplasmic reticulum. The observed increase in the number of scalloped sacs usually is accompanied by a significant expansion of the whole system of endoplasmic reticulum. Some of the vacuoles seen light-microscopically must indeed be expanded cisternae of endoplasmic reticulum. The findings are discussed in light of the possibility that the scalloped sacs and the endoplasmic reticulum may be involved in active transport of sodium ions.     

Evidence,information, and surprise

A numerical measure for evidence is defined in a probabilistic framework. The established mathematical concept of information or entropy (as defined in ergodic theory) can be obtained from this definition in a special case, although in general information is greater than evidence. In another, somewhat complementary, special case a numerical measure for surprise is derived from the definition of evidence. Some applications of the new concept of evidence are discussed, concerning statistics in general and the special kind of statistics performed by neurophysiologists, when they analyze the response of neurons, and perhaps by the neurons themselves.     

Protoplasmic streaming in the giant unicellular green algaAcetabularia mediterranea

Summary Protoplasmic Streaming inAcetabularia mediteranea has been studied by microcinematography in 1. germinating zygotes, 2. germlings before the differentiation of rhizoids and apices, 3. young cells with rhizoids and apices, 4. vegetative cells-several centimeters in length, 5. cells with a maximum sized cap, containing secondary nuclei, and 6. cells after cyst formation. Intracellular transport is found to occur at a network-system of thin filaments and at a different system of headed streaming bands. At the network of filaments chloroplasts are found to move at a velocity of 1–2 m/sec. Headed streaming bands move along the filaments and may lead without interruption from the rhizoid to the apex of the cell andvice versa. The front zone of the streaming bands is occupied by a leading cytoplasmic head-structure. Small vesicles, polyphosphate granula and secondary nuclei are the predominant moving structures in headed streaming bands. The velocity of these particles is found to be 3–11 m/sec. The filament system is found during all developmental stages. Headed streaming bands are undetectable in germinating zygotes and develop from small cytoplasmic droplets in germlings to broad heavily loaded bands in the huge vegetative cell.Transport of secondary nuclei by headed streaming bands is not observed during mitotic divisions and after cyst formation, though moving bands are still present for several weeks after cyst formation.     

Typology Reconsidered: Two Doctrines on the History of Evolutionary Biology

Recent historiography of 19th century biology supports the revision of two traditional doctrines about the history of biology. First, the most important and widespread biological debate around the time of Darwin was not evolution versus creation, but biological functionalism versus structuralism. Second, the idealist and typological structuralist theories of the time were not particularly anti-evolutionary. Typological theories provided argumentation and evidence that was crucial to the refutation of Natural Theological creationism. The contrast between functionalist and structuralist approaches to biology continues today, and the historical misunderstanding of 19th century typological biology may be one of its effects. This historical case can shed light on current controversies regarding the relevance of developmental biology to evolution.