On a possible relationship between bacterial endospore formation and the origin of eukaryotic cells

The acquisition of intracellular organelles, including mitochondria and plastids and a membrane-bounded nucleus, have been postulated to be key events in the development of the eukaryotic from the prokaryotic ancestral cell. The two major hypotheses to account for such acquisitions are: (1) primitive cells originally obtained organelles by engulfing free-living prokaryotes which then entered into symbiotic association (“endosymbiosis”) with them; (2) organelles arose through the engulfment by the primitive cell of part of its own cytoplasm. To some extent, the former hypothesis has received most support, because endosymbiosis is known to occur in extant organisms, whilst the latter hypothesis has received less support, because cytoplasmic engulfment by prokaryotes is not now thought to occur. However, during the process of endospore formation by extant bacteria, the protoplast within the single cell is observed to divide in a unique manner such that the cell in effect engulfs a portion of its own cytoplasm. The process is strikingly similar to the engulfment suggested by the second hypothesis to have initiated the evolution of eukaryotes. The engulfed cytoplasm is bounded by a double membrane within the “mother cell” and contains enzymes, ribosomes and a complete genome. In many respects this parallels the supposed primitive eukaryotic state and, it is argued, confers potential advantages on the cell, particularly through the control that the “mother cell” can exert on the enclosed compartment. It is hypothesized that bacterial endospore formation is therefore one product of evolution from an early engulfment event that led also to the development of complex eukaryotic cells.     

pH and expansin action on single suspension-cultured tomato (Lycopersicon esculentum) cells

The aim of this study was to measure key material properties of the cell walls of single suspension-cultured plant cells and relate these to cell-wall biochemistry. To this end, micromanipulation was used to compress single tomato cells between two flat surfaces until they ruptured, and force-deformation data were obtained. In addition to measuring the bursting force, we also determined the elastic (Young’s) modulus of the cell walls by matching low strain (≤20% deformation) experimental data with a cell compression model, assuming linear elastic cell walls. The walls were most elastic at pH 4.5, the pH optimum for expansin activity, with an elastic modulus of 2.0 ± 0.1 GPa. Following the addition of exogenous expansins, cell walls became more elastic at all pH values. Western blot analysis of proteins from walls of cultured cells revealed the presence of expansin epitopes, suggesting that the inherent pH dependence of elasticity and other compression phenomena is related to the presence of endogenous expansin proteins and their wall-loosening ability. Although strict application of the linear-elastic model could not be applied to large deformations—for example, up to cell bursting—because of irreversible behaviour, the deviation of the data from the model was generally small enough to allow estimation of the strain in the cell wall at failure. This strain was greater at pH 4.5 and when expansins were added to the suspension. The changes in elasticity are consistent with suggestions about the mode of expansin action. The estimated strains at failure are compatible with data on the failure of Acetobacter-derived cellulose–xyloglucan composites and proposed mechanisms of such failure. Through the measurement of cell-wall material properties using micromanipulation, it may be possible to understand more fully how cell-wall composition, structure and biochemistry lead to cell mechanical behaviour.     

The evolution of male-female dimorphism: Older than sex?

This contribution considers the evolution of a dimorphism with respect to cell fusion characteristics in a population of primitive cells. These cells reproduce exclusively asexually. The evolution towards asymmetric fusion behaviour of cells is driven by selection promoting horizontal transfer of an endosymbiontic replicator. It is concluded that evolution of asymmetric cell fusion in this scenario is more likely than evolution of sexual differentiation in a sexually reproducing population. Pre-existing dimorphism with respect to cell fusion may thus have been the basis for the establishment of sexual differentiation at the level of gamete fusion, and this in turn is fundamental to the evolution of two different sexes, male and female.     

Nuclear matrix of the most primitive eukaryote Archezoa

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The Acoela: on their kind and kinships, especially with nemertodermatids and xenoturbellids (Bilateria incertae sedis)

Acoels are among the simplest worms and therefore have often been pivotal in discussions of the origin of the Bilateria. Initially thought primitive because of their “planula-like” morphology, including their lumenless digestive system, they were subsequently dismissed by many morphologists as a specialized clade of the Platyhelminthes. However, since molecular phylogenies placed them outside the Platyhelminthes and outside all other phyla at the base of the Bilateria, they became the focus of renewed debate and research. We review what is currently known of acoels, including information regarding their morphology, development, systematics, and phylogenetic relationships, and put some of these topics in a historical perspective to show how the application of new methods contributed to the progress in understanding these animals. Taking all available data into consideration, clear-cut conclusions cannot be made; however, in our view it becomes successively clearer that acoelomorphs are a “basal” but “divergent” branch of the Bilateria.     

Control and Regulatory Mechanisms Associated with Thermogenesis in Flying Insects and Birds

Most insects and birds are able to fly. The chitin made exoskeleton of insects poses them several constraints, and this is one the reasons they are in general small sized animals. On the other hand, because birds possess an endoskeleton made of bones they may grow much larger when compared to insects. The two taxa are quite different with regards to their general “design” platform, in particular with respect to their respiratory and circulatory systems. However, because they fly, they may share in common several traits, namely those associated with the control and regulatory mechanisms governing thermogenesis. High core temperatures are essential for animal flight irrespective of the taxa they belong to. Birds and insects have thus evolved mechanisms which allowed them to control and regulate high rates of heat fluxes. This article discusses possible convergent thermogenic control and regulatory mechanisms associated with flight in insects and birds.     

Cytogenetic study of three edible chrysanthemum cultivars

During past decades, edible chrysanthemum (Dendranthema × grandiflorum) has become a popular dietary supplement. However, only numberable cultivars are available, cytogenetic studies especially meiotic behaviour in them remaind largely unexplored. In the present study, we analyzed the karyotype and meiotic behaviour during microsporogenesis in accessions of edible chrysanthemum. This information can be useful in cultivar improvement, as well as in elucidation of the evolution of the species. The three edible cultivars studied all had an intermediate type of interphase nucleus, and their mitotic prophase chromosomes were of the interstitial type. The chromosome number of “Baohuatangyijin”(Bn) varies from 55 to 62. “Zifengmudan”(Zn) has the karyotype 2n = 54 = 38m + 12sm + 4st, while “Jingxingxiying”(Jg) has 2n = 55 = 38m + 15sm + 2st. Male meiosis was largely normal. Mean pairing configuration of Zn at diakinesis and metaphase I was 0.94I + 25.07II + 0.14III + 0.63IV, while for Jg, the equivalent was 1.32I + 24.64II + 0.16III + 0.85IV + 0.05V + 0.04VI. Chromatid separation was normal during anaphase I and anaphase II in most meiocytes. All three edible chrysanthemums appear to be allopolyploid, and the edible type is probably more primitive than the ornamental type. However, the edible type is probably more closely related to the ornamental than to the medicinal type.     

Edward Hitchcock’s Pre-Darwinian (1840) “Tree of Life”

The “tree of life” iconography, representing the history of life, dates from at least the latter half of the 18th century, but evolution as the mechanism providing this bifurcating history of life did not appear until the early 19th century. There was also a shift from the straight line, scala naturae view of change in nature to a more bifurcating or tree-like view. Throughout the 19th century authors presented tree-like diagrams, some regarding the Deity as the mechanism of change while others argued for evolution. Straight-line or anagenetic evolution and bifurcating or cladogenetic evolution are known in biology today, but are often misrepresented in popular culture, especially with anagenesis being confounded with scala naturae. Although well known in the mid 19th century, the geologist Edward Hitchcock has been forgotten as an early, if not the first author to publish a paleontologically based “tree of life” beginning in 1840 in the first edition of his popular general geology text Elementary Geology. At least 31 editions were published and those between 1840 and 1859 had this “paleontological chart” showing two trees, one for fossil and living plants and another for animals set within a context of geological time. Although the chart did not vary in later editions, the text explaining the chart did change to reflect newer ideas in paleontology and geology. Whereas Lamarck, Chambers, Bronn, Darwin, and Haeckel saw some form of transmutation as the mechanism that created their “trees of life,” Hitchcock, like his contemporaries Agassiz and Miller, who also produced “trees of life,” saw a deity as the agent of change. Through each edition of his book Hitchcock denounced the newer transmutationist hypotheses of Lamarck, then Chambers, and finally Darwin in an 1860 edition that no longer presented his tree-like “paleontological chart.”     

The hypoblast (visceral endoderm): an evo-devo perspective

When amniotes appeared during evolution, embryos freed themselves from intracellular nutrition; development slowed, the mid-blastula transition was lost and maternal components became less important for polarity. Extra-embryonic tissues emerged to provide nutrition and other innovations. One such tissue, the hypoblast (visceral endoderm in mouse), acquired a role in fixing the body plan: it controls epiblast cell movements leading to primitive streak formation, generating bilateral symmetry. It also transiently induces expression of pre-neural markers in the epiblast, which also contributes to delay streak formation. After gastrulation, the hypoblast might protect prospective forebrain cells from caudalizing signals. These functions separate mesendodermal and neuroectodermal domains by protecting cells against being caught up in the movements of gastrulation.     

The accumulation of basic nonhemoglobin proteins during the differentiation of primitive (embryonic) red cells of amphibia

The preferential accumulation of hemoglobin is a characteristic of the differentiation of definitive (adult) red blood cells. Since primitive (embryonic or larval) red blood cells have many properties which contrast with definitive red cells, the accumulation of red cell proteins was analyzed during the differentiation of primtive red cells to determine whether or not hemoglobin was the only protein which showed a substantial increase in amount. Primitive red cells of amphibia were used because the mature circulating cell retains large amounts (13%) of specific, well characterized, nonhemoglobin proteins (CP). Preparations of primitive red cells enriched in immature cells were obtained from the circulation of bullfrog tadpoles recovering from phenylhydrazine-induced anemia. The amount of CP, determined by electrophoresis, imunodiffusion, and ion-exchange chromatography, was compared for red cells from normal animals and anemic animals. Mature cells contained three to six times the amount of CP and three times the amount of hemoglobin found in the population of red cells enriched in immature cells. The accumulation of CP during maturation of primitive red cells indicates that differentiation of primitive red cells is less restrictive than differentiation of definitive red cells. Since the primitive red cell is less specialized than the definitive red cell, it is possible that primitive red cells have several roles in the developing animal, in contrast to the single role of synthesizing and maintaining hemoglobin in the adult animal.     

An ECM substratum allows mouse mesodermal cells isolated from the primitive streak to exhibit motility similar to that inside the embryo and reveals a deficiency in the T/T mutant cells

The mesodermal cell layer is created by ingression and migration of the cells from the primitive streak region in mouse embryos on day 7 of pregnancy. In order to study the mechanisms of mesodermal cell migration during development, the mesodermal cells isolated from the primitive streak were cultured on various substrata, and cell behaviour and motility were analysed with a time-lapse video system. The mesodermal cells on the surface of extracellular matrix (ECM)-coated dishes (ECM produced by bovine corneal endothelial cells) showed extensive migration at a mean rate of approx. 50 micron h-1. They also showed frequent cell division and exhibited contact paralysis of lamellipodia and contact inhibition of movement. On plastic or glass surfaces, however, the mesodermal cells became more flattened and less motile (approx. 20-30 micron h-1). Cell shape and mean rate of movement on the ECM were very similar to those in situ, as investigated in a previous study (Nakatsuji, Snow & Wylie, 1986). Therefore, this culture condition could provide a useful experimental system for analysing the cellular basis of normal and abnormal morphogenetic movements in mouse embryos. Employing such a culture system, we studied motility of the mesodermal cells from embryos homozygous for Brachyury (T) mutation, which are lethal at the midgestation stage in utero. Histological observations have suggested that anomalous morphogenesis of the T/T embryos may be brought about by defects in migration of the mesodermal cells derived from the primitive streak. When mesodermal cells from the primitive streak of the T/T mutant embryos on days 8-9 were cultured on the ECM substratum, mean rate of cell migration was significantly reduced compared to cells from normal embryos. Results support the idea of retarded migration by the mutant mesodermal cells as an important factor causing abnormalities in morphogenesis.     

Nuclear matrix of the most primitive eukaryote Archezoa

Accumulating evidence suggests that unicellularArchezoa are the most primitive eukaryotes and their nuclei are of significance to the study of evolution of the eukaryotic nucleus. Nuclear matrix is an ubiquitous important structure of eukaryotic nucleus; its evolution is certainly one of the most important parts of the evolution of nucleus. To study the evolution of nuclear matrix, nuclear matrices ofArchezoa are investigated.Giardia lamblia cells are extracted sequentially. Both embedment-free section EM and whole mount cell EM of the extracted cells show that, like higher eukaryotes, this species has a residual nuclear matrix in its nucleus and rich intermediate filaments in its cytoplasm, and the two networks connect with each other to form a united network. But its nuclear matrix does not have nucleolar matrix and its lamina is not as typical as that of higher eukaryotes; Western blotting shows that lamina ofGiardia and two otherArchezoa Entanzoeba invadens andTrichomonas vaginali all contain only one polypeptide each which reacts with a mammalia anti-lamin polyclonal serum and is similar to lamin B (67 ku) of rnammiia in molecular weight. According to the results and references, it is suggested that nuclear matrix is an early acquisition of the eukaryotic nucleus, and it and the “eukaryotic chromatin” as a whole must have originated very early in the process of evolution of eukaryotic cell, and their origin should be an important prerequisite of the origin of eukaryotic nucleus: in the lamin (gene) family, B-type lamins (gene) should be the ancestral typz and that A-type lamins (gene) might derive therefrom. Project supported by the National Natural Science Foundation of China (Grant No. 3870254).     

Six major steps in animal evolution: are we derived sponge larvae?

A review of the old and new literature on animal morphology/embryology and molecular studies has led me to the following scenario for the early evolution of the metazoans. The metazoan ancestor, "choanoblastaea," was a pelagic sphere consisting of choanocytes. The evolution of multicellularity enabled division of labor between cells, and an "advanced choanoblastaea" consisted of choanocytes and nonfeeding cells. Polarity became established, and an adult, sessile stage developed. Choanocytes of the upper side became arranged in a groove with the cilia pumping water along the groove. Cells overarched the groove so that a choanocyte chamber was formed, establishing the body plan of an adult sponge; the pelagic larval stage was retained but became lecithotrophic. The sponges radiated into monophyletic Silicea, Calcarea, and Homoscleromorpha. Homoscleromorph larvae show cell layers resembling true, sealed epithelia. A homoscleromorph-like larva developed an archenteron, and the sealed epithelium made extracellular digestion possible in this isolated space. This larva became sexually mature, and the adult sponge-stage was abandoned in an extreme progenesis. This eumetazoan ancestor, "gastraea," corresponds to Haeckel's gastraea. Trichoplax represents this stage, but with the blastopore spread out so that the endoderm has become the underside of the creeping animal. Another lineage developed a nervous system; this "neurogastraea" is the ancestor of the Neuralia. Cnidarians have retained this organization, whereas the Triploblastica (Ctenophora+Bilateria), have developed the mesoderm. The bilaterians developed bilaterality in a primitive form in the Acoelomorpha and in an advanced form with tubular gut and long Hox cluster in the Eubilateria (Protostomia+Deuterostomia). It is indicated that the major evolutionary steps are the result of suites of existing genes becoming co-opted into new networks that specify new structures. The evolution of the eumetazoan ancestor from a progenetic homoscleromorph larva implies that we, as well as all the other eumetazoans, are derived sponge larvae.     

“Theory” in Theory and Practice

A central obstacle to accepting evolution, both among students and the general public, is the idea that evolution is “just a theory,” where “theory” is understood in a pejorative sense as something conjectural or speculative. Although scientists and textbooks constantly explain that the scientific use of “theory” is quite different, the pejorative use continues to cause confusion, in part because of its deep roots in a popular, Baconian, understanding of science. A constructivist approach, whereby students are helped to examine the adequacy of their preconceptions about “theory” for themselves and to revise or replace them appropriately, is recommended.     

“Silenced” polydendrocytes: a new cell type within the oligodendrocyte progenitor cell population?

Oligodendrocyte progenitor cells (OPCs) were first described more than two decades ago. Novel labeling techniques have shown them to be cells with more than just progenitor functions, with their classification as a fourth glial cell type in addition to astrocytes, oligodendrocytes, and microglial cells. Another term used for this cell type is polydendrocytes, owing to both their morphology and to the evolving knowledge about their diverse functions. Recently, an exclusive hallmark of neurons—the generation of action potentials—became debatable, because a subset of polydendrocytes was reported to generate action potentials in response to adequate stimuli. The new technique of inducible reporter gene expression has brought new insights into the fate and function of polydendrocytes. In recent studies, so-called “silenced” OPCs were detected in cortical tissue, and which underwent proliferation with subsequent cell cycle exit, but without any signs of differentiation. Within this review, we focus on the identification of this new subset of polydendrocytes and their possible functions within cortical networks.     

The Evolution of Bioluminescent Oxygen Consumption as an Ancient Oxygen Detoxification Mechanism

Endogenous reductants such as hydrogen sulfide and alkylthiols provided free radical scavenging systems during the early evolution of life. The development of oxygenic photosynthesis spectacularly increased oxygen levels, and ancient life forms were obliged to develop additional antioxidative systems. We develop here the hypothesis of how ``prototypical' bioluminescent reactions had a plausible role as an ancient defense against oxygen toxicity through their ``futile' consumption of oxygen. As oxygen concentrations increased, sufficient light would have been emitted from such systems for detection by primitive photosensors, and evolutionary pressures could then act upon the light emitting characteristics of such systems independently of their use as futile consumers of oxygen. Finally, an example of survival of this ancient mechanism in present-day bioluminescent bacteria (in the Euprymna scolopes–Vibrio fischeri mutualism) is discussed. Once increasing ambient oxygen levels reached sufficiently high levels, the use of ``futile' oxygen consumption became too bioenergetically costly, so that from this time the evolution of bioluminescence via this role was made impossible, and other mechanisms must be developed to account for the evolution of bioluminescence by a wide range of organisms that patently occurred after this (e.g., by insects). Received: 25 May 2000 / Accepted: 14 November 2000     

Identification Keys,the “Natural Method,” and the Development of Plant Identification Manuals

The origins of field guides and other plant identification manuals have been poorly understood until now because little attention has been paid to 18th century botanical identification guides. Identification manuals came to have the format we continue to use today when botanical instructors in post-Revolutionary France combined identification keys (step-wise analyses focusing on distinctions between plants) with the “natural method” (clustering of similar plants, allowing for identification by gestalt) and alphabetical indexes. Botanical works featuring multiple but linked techniques to enable plant identification became very popular in France by the first decade of the 19th century. British botanists, however, continued to use Linnaeus’s sexual system almost exclusively for another two decades. Their reluctance to use other methods or systems of classification can be attributed to a culture suspicious of innovation, anti-French sentiment and the association of all things Linnaean with English national pride, fostered in particular by the President of the Linnean Society of London, Sir James Edward Smith. The British aversion to using multiple plant identification technologies in one text also helps explain why it took so long for English botanists to adopt the natural method, even after several Englishmen had tried to introduce it to their country. Historians of ornithology emphasize that the popularity of ornithological guides in the 19th and 20th centuries stems from their illustrations, illustrations made possible by printing technologies that improved illustration quality and reduced costs. Though illustrations are the most obvious features of late 19th century and 20th century guides, the organizational principles that make them functional as identification devices come from techniques developed in botanical works in the 18th century.     

细胞核起源研究的意义和研究途径的探讨

细胞核的起源是真核细胞进化形成的关键。回顾了过去几十年国内外对细胞核起源问题的探索历程,通过多年的摸索找到了一个条切实可行的探索细胞核起源问题的途径。其要点：在一系列的进化环节中首先抓住原始性的细胞核这一重要环节,探明原始性细胞核的特性,解决了从原始核到典型细胞核的进化问题,原始性细胞核自身的起源问题也就有了基础,为探源始性细胞核的特性,需要在现存的原生生物中间寻找最原始的类群,然后对它们的细胞核进行尽可能深入地和多方面地研究,对所得结果作进化地分析,以期提出一个原始性细胞核的模型,依据这个模型也就可对典型的细胞核的进化形成和原始核自身的起源作出推论,而这个原始性细胞核的模型,依据这个模型也就可以对典型细胞核的进化形成和原始核自身的起源作出推论,这些推论是可以设法加以检验的,不仅可以检验这些推论的正确性,而且对原始核模型的建立是重要的,可以据之加以发展,修正,甚至否定,沿此途径已经否定了原始性细胞核的涡鞭毛虫核模型,进而提出了双滴虫核模型。     

Animal tool-use

The sight of an animal making and using a tool captivates scientists and laymen alike, perhaps because it forces us to question some of our ideas about human uniqueness. Does the animal know how the tool works? Did it anticipate the need for the tool and make it in advance? To some, this fascination with tools seems arbitrary and anthropocentric; after all, animals engage in many other complex activities, like nest building, and we know that complex behaviour need not be cognitively demanding. But tool-using behaviour can also provide a powerful window into the minds of living animals, and help us to learn what capacities we share with them - and what might have changed to allow for the incontrovertibly unique levels of technology shown by modern humans.     

NEMERTINES AS POSSIBLE ANCESTORS OF THE VERTEBRATES

1. Primitive vertebrates have many characters which do not appear to be immediately represented in any obvious precursor. 2. The hypothesis is put forward that creeping, or bottom-living worms, classifiable as nemertines, attempted to recolonize the water by acquiring buoyancy, by swimming or both. 3. Some of those that succeeded were able to do so by adopting a method of filter-feeding made adequate by the development of gill slits, and then converting the proboscis rudiment into a semi-rigid structure (the notochord) essential for oscillatory swimming movements. These were the precursors of the protochordates and the vertebrates. 4. The acquisition of the ability to swim efficiently depended on the simultaneous development of distance receptors and the co-ordination of the information that they provided with that provided by proprioceptors on the one hand and with the changing motor systems on the other. This entailed the development of a far more extensive nervous system, probably by incorporating into it much more of the dorsal ectoderm, and superimposing a primarily sensori-motor system on to the more vegetative system already present in nemertines. This was achieved by elaboration of the placodal folding of the ectoderm which is characteristic of many nemertine embryos of the present day. In this manner a new central nervous system was combined and integrated with the existing more primitive ‘autonomic’ system and the cephalic ganglia of the nemertines became incorporated in the hypothalamus. 5. The hypothesis suggests likely precursor tissues from which the following vertebrate structures could plausibly have developed: olfactory organ, lateral line system, anterior and posterior pituitary, thyroid, pineal organ, chloride-secreting cells, oxyntic cells, notochord, urinogenital system, liver, and several others. 6. The possible origins of the vertebrate eye, the somitic muscles, and some other fundamental features of the vertebrates are discussed. The evidence is still equivocal. 7. The acquisition of filter-feeding and swimming by nemertine worms would be expected to bring into operation a whole new system for the selection of appropriate genes and thus inaugurate a period of very rapid and probably divergent evolution, made even more rapid by the enormous advantages gained by animals acquiring efficient form-vision together with rapid and well co-ordinated movements. It is thought likely that the rapid evolution of this group of animals, which possessed few features conducive to clear fossil remains, accounts for the apparent break in the continuity of the story of animal evolution which has until now made the origin of the vertebrates such an enigma. 8. It is suggested that studies of the proteins and secretions of the various organs of the nemertines and their comparison with those of the organs of the more primitive vertebrates might lead to the establishment of some important homologies.