THE SYSTEMATICS AND PHYLOGENETIC RELATIONSHIPS OF VETULICOLIANS

Abstract:  Vetulicolians have variously been considered to be unusual arthropods, stem-group deuterostomes or relatives of the tunicates. They are known from a number of Cambrian Lagerstätten, and are particularly diverse in the Chengjiang biota of Yunnan Province, China. We recognize two classes, Vetulicolida and Banffozoa, which together form a monophyletic group. Within the Chinese collections we also identify two new species and recognize one new genus: Vetulicola monile sp. nov. and Bullivetula variola gen. et sp. nov. The evidence from new and previously described specimens is used to undertake a phylogenetic analysis and to evaluate a range of hypotheses for the affinities of vetulicolians. Given the difficulties of interpreting features in enigmatic fossils and the apparently contradictory set of characters possessed by vetulicolians, it is not possible on current evidence to reach an unequivocal conclusion regarding the phylogenetic position of the group. One possibility is that they are a sister group of arthropods that lost limbs but gained gill structures analogous to those of deuterostomes, but several features remain unexplained by this model. If they are protostomes, a more generally parsimonious position is close to the kinorhynchs. An alternative is that they are deuterostomes, although a placement at the base of the clade is not supported by the evidence. If they are deuterostomes, it is more likely that they are close to the tunicates.     

The earliest history of the deuterostomes: the importance of the Chengjiang Fossil-Lagerst?tte

While the broad framework of deuterostome evolution is now clear, the remarkable diversity of extant forms within this group has rendered the nature of the ancestral types problematic: what, for example, does the common ancestor of a sea urchin and lamprey actually look like? The answer to such questions can be addressed on the basis of remarkably well-preserved fossils from Cambrian Lagerstätten, not least the celebrated Chengjiang Lagerstätte (Yunnan, China). This deposit is particularly important because of its rich diversity of deuterostomes. These include some of the earliest known representatives, among which are the first vertebrates, as well as more enigmatic groups, notably the vetulicolians and yunnanozoans. The latter groups, in particular, have been the subject of some radical divergences in opinion as to their exact phylogenetic placements. Here, we both review the known diversity of Chengjiang deuterostomes and in particular argue that the vetulicolians and yunnanozoans represent very primitive deuterostomes. Moreover, in the latter case we present new data to indicate that the yunnanozoans are unlikely to be any sort of chordate.     

昆明地区下寒武统沧浪铺组古虫动物新发现

文中详细描述昆明地区下寒武统沧浪铺组乌龙箐段古虫动物新种Vetulicola longbaoshanensis sp.nov.,并与相关属种作比较和讨论。古虫动物主要产于下寒武统筇竹寺组玉案山段的澄江动物群中,沧浪铺组乌龙箐段的地质年代较澄江动物群晚,该发现对研究古虫动物的演化具有重要意义。古虫动物的分类位置至今难以确定,争论的焦点多集中在古虫动物前体鳃囊的构造解释上。新材料鳃囊保存得完整清晰,为研究古虫动物的分类位置提供了新证据。     

Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte,North Greenland,and the polarity of morphological characters in basal deuterostomes

Abstract: Vetulicolians are problematic Cambrian fossils with a debated phylogenetic history. Here, we describe two vetulicolian specimens from the Lower Cambrian Sirius Passet locality in North Greenland. One of the specimens is assigned to Ooedigera peeli gen. et sp. nov, whereas the other is retained under open nomenclature. The mode of tail flexibility has been debated in the literature, and we argue here that the tail normally flexed laterally to generate power strokes rather than dorsoventrally. The phylogenetic relationships of vetulicolians are discussed in the light of current knowledge of deuterostome phylogeny and morphology, and it is concluded that the best hypothesis on currently available evidence is that vetulicolians are a clade or paraphyletic assemblage of stem‐Deuterostomata. The presence of a voluminous, sediment‐filled anterior chamber suggests that the atrium may be a synapomorphy of deuterostomes.     

Vetulicolians—are they deuterostomes? chordates?

A recent paper by Shu et al.(1) reinterprets the fossil Vetulicola and related forms, all from the Lower Cambrian, as basal deuterostomes, assigning them their own phylum, Vetulicolia. Their conclusion is based on the presence of structures resembling gill slits and a trunk-like region that shows evidence of segmentation. This report summarizes the fossil evidence for their interpretation and evaluates a possible alternative, that vetulicolians may instead be tunicate-like chordates. Implications for our understanding of the nature of the primitive deuterostome (and chordate) body plan are discussed. .     

用红外方法对蝇翅视动行为的探索

本文报告了利用红外装置对蝇翅视动行为实验研究的初步结果及其分析:1.在红外探测器探测到的信号中找到了一个能反映蝇翅拍动幅度的参数.2.双侧、单侧刺激域的宽度及刺激域的高度对视动反应发生几率在一定范围内正相关,当超过一阈值(即饱和阈值)后,即出现稳定的视动反应,它们的饱和阈值分别为60°,30°,40°刺激条纹的亮度生有类似情况.刺激条纹的运动速度在一定范围内对视动反应无影响.3.当刺激没有达到饱和时,蝇翅出现断续的典型的视动反应,即“0-1波动反应”.4.单侧条纹由前向后运动时,蝇翅出现典型反应,而条纹从后向前运动时,不出现典型的视动反应或反应很弱.双侧刺激时,条纹向前运动几乎不诱发反应;条纹向后运动诱发明显的蝇翅视动反应,且蝇翅平面的方向在拍动过程中发生变化.     

CELL POPULATION KINETICS OF THE SCLEROBLAST DURING ETHISTERONE-INDUCED ANAL-FIN PROCESS FORMATION IN ADULT FEMALES OF THE MEDAKA, ORYZIAS LATIPES

Proliferation and migration of the scleroblasts and their precursor cells during ethisterone-induced anal-fin process formation in adult females of the medaka, Oryzias latipes , were investigated using a radioautographic technique with ³H-thymidine. In the posterior margin of the joint plate, rapid increases in labeling indexes and scleroblast population began simultaneously in the central portion on the third day of ethisterone treatment, and the population became approximately ten times that in the control fish on the fifth day. However, nomarked increase in scleroblast population occurred in the peripheral portion, although the scleroblasts showed nearly the same labeling index as those in the central portion throughout the experiment. These observations indicate that the scleroblasts and their precursor cells migrated from the peripheral to the central portion along the proxi-distal axis of the joint plate. Some cells in the inner and hind spaces also migrated to the posterior margins and subsequently developed into scleroblasts. It may be concluded, therefore, that the scleroblasts are derived from precursor cells situated not only along the posterior margins but also in the adjacent spaces. These precursor cells proliferate and/or migrate to the central portion of the posterior margins and aggregate to form the scleroblast mass.     

AXONAL TRANSPORT OF PROTEINS IN THE HYPOTHALAMO-NEUROHYPOPHYSIAL SYSTEM OF THE RAT

—Axonal transport of proteins in the hypothalamo-neurohypophysial system of the rat was studied after a local injection of [³⁵S]cysteine in the region of the supraoptic nucleus. The migration of labelled proteins was followed by measuring the specific radioactivity of the proteins in various parts of the hypothalamo-neurohypophysial tract. Between 2 and 4 h after the isotope injection there was a sharp increase in the protein-bound specific radioactivity of the posterior pituitary lobe, demonstrating that a transport of ³⁵S-labelled proteins had occurred from the supraoptic nucleus to the neurohypophysis. The rate of the transport was 2-3 mm/h. During the first 24 h after the injection a continuous accumulation of labelled material occurred in the neural lobe. Considerable radioactivity could still be recovered 6 days after the isotope injection. Fractionation of the neurohypophysial proteins by polyacrylamide gel electrophoresis revealed that approximately 90 per cent of the radioactivity of the soluble proteins was recovered in a single protein fraction. Labelling of this fraction was not observed until 2 h after isotope injection. The radioactivity increased markedly up to 4 h. It is suggested that this protein component is involved in the neurohypophysial response to osmotic stress since the protein disappeared from the posterior lobe upon dehydration of the rat.     

湘西北志留纪胴甲鱼化石

本文记述了采自湘西北澧县石门水库附近秀山组上段隶属云南鱼目的 Shimenolepis graniferus gen. et sp. nov. 和 QuJinolepidae gen. et sp. indet. 化石,对含鱼层的时代也进行了讨论.     

CYTODIFFERENTIATION IN THE ROSY MUTANT OF DROSOPHILA MELANOGASTER

In the rosy mutant of Drosophila melanogaster, two types of autofluorescent cytoplasmic inclusions are found in the cells of the posterior region of the fatbody at the prepupal stage. Bright yellow autofluorescent granules accumulating within larger inclusions clearly demarcate this area of the fatbody which also contains cobalt blue fluorescent globular material. Such inclusions were not noted in the normal Ore-R strain at this stage nor in the series of mutant strains examined other than the rosy² and maroon-like mutants. The pattern of biochemical deviation of the latter two mutants is known to be identical to that of the rosy mutant, and a portion of this mutant upset can be ascribed to the absence of xanthine dehydrogenase. These mutants lack the products of enzyme activity, uric acid and isoxanthopterin, and accumulate their precursors, hypoxanthine and 2-amino-4-hydroxypteridine. Chromatographic studies on the fatbody of rosy prepupae have shown that 2-amino-4-hydroxypteridine is limited to the posterior region; this correspondence in location as well as color of fluorescence indicates that the cobalt blue auto fluorescent globules in the fatbody contain 2-amino-4-hydroxypteridine. In the normal strain, isoxanthopterin was identified in the chromatograms of the posterior region of the fatbody, but it was not obtained from the anterior region of the fatbody. On the other hand, xanthine dehydrogenase activity could be demonstrated throughout the fatbody of the normal strain. The restriction of isoxanthopterin to a certain group of fat cells in the wild type strain and its absence from other fat cells can be explained by the differential distribution of its immediate precursor, 2-amino-4-hydroxypteridine, as displayed in the mutant rosy.     

大黄鱼精子的超微结构

大黄鱼的精子由头产和尾部两部分组成。头部结构较为独特,其腹侧有一较大的细胞核,背部有中心粒复合体。头部的后端是袖套。细胞核的腹面稍向外突出背面则稍向内凹。细胞核中的染以质浓缩成致密的团块状。团块状的染色质之间分布着松散的纤维状染色质。植入窝位于细胞核的背部表面,由细胞核背面向内凹陷而成,呈一沟状,其走向与精子的长轴平行。     

FINE STRUCTURE OF THE MUCOCYSTS OF COLACIUM CALVUM (EUGLENOPHYCEAE)1

The characteristic stalk of Colacium calvum Stein is formed from carbohydrate extruded by granules in the anterior portion of the cell. These granules exhibit the characteristics of mucocysts and their contents have two electron densities. A narrow extension of each granule extends past neighboring muciferous bodies to attachment sites in the posterior notch of the pellicle strip. The granule contains neutral polysaccharide which resembles that in the peripheral cisternae of the Golgi apparatus. The attachment site is structurally complex.     

STUDIES ON THE FINE STRUCTURE OF THE MAMMALIAN TESTIS : I. DIFFERENTIATION OF THE SPERMATIDS IN THE CAT (FELIS DOMESTICA)

The differentiation of cat spermatids was studied in thin sections examined with the electron microscope. The Golgi complex of the spermatid consists of a central aggregation of minute vacuoles, partially surrounded by a lamellar arrangement of flattened vesicles. In the formation of the acrosome, one or more moderately dense homogeneous granules arise within vacuoles of the Golgi complex. The coalescence of these vacuoles and their contained granules gives rise to a single acrosomal granule within a sizable membrane-limited vacuole, termed the acrosomal vesicle. This adheres to the nuclear membrane and later becomes closely applied to the anterior two-thirds of the elongating nucleus to form a closed bilaminar head cap. The substance of the acrosomal granule occupies the narrow cleft between the membranous layers of the cap. The caudal sheath is comprised of many straight filaments extending backward from a ring which encircles the nucleus at the posterior margin of the head cap. Attention is directed to the frequent occurrence of pairs of spermatids joined by a protoplasmic bridge and the origin and possible significance of this relationship are discussed.     

THE NEUROENDOCRINE SYSTEM AND STOMATOGASTRIC NERVOUS SYSTEM OF THE ADULT TSETSE FLY GLOSSINA MORSITANS

The brain of Glossina morsitans Westwood contains four groups of neurosecretory cells which are stainable with chrome haematozylin and phloxin. The axons of these cells form a pair of nervi corporis cardiaci which pass posteriorly from the brain and innervate the corpora cardiaca and corpus allatum before uniting with a small ganglion posterior to the corpora cardiaca. This ganglion is considered to represent the fusion of the fusion of the hypocerebral and ventricular ganglia which remain separate in other insects.
There is no frontal ganglion in the adult Glossina and the recurrent nerve fuses with one of the nervi corporis cardiaci immediately behind the brain. The oesophageal nerves arising from the fused hypocerebral and ventricular ganglia innervate the oesophagus in the anterior part of the thorax, the proventriculus and the posterior extension of the oesophagus close to the crop. These nerves possess both sensory and motor nerve endings. The differences which exist between Glossina and other cyclorrhaphous Diptera with respect to their neuroendocrine/stomatogastric system are noted and considered in terms of the control of neuroendocrine function.     

四川内江晚侏罗世中国龟科一新属

描述了产自四川内江晚侏罗世的中国龟科（Ｓｉｎｅｍｙｄｉｄａｅ）化石一新属新种──叶氏香港龟Ｈｏｎｇｋｏｎｇｏｃｈｅｌｙｓ ｙｅｈｉ。它以椎盾较横宽、第７、８椎板较退化、上臀板缺失、具腹甲后窗等特征区别于中国龟属（Ｓｉｎｅｍｙｓ）和满洲龟属（Ｍａｎｃｈｕｒｏｃｈｅｌｙｓ）。这是中国龟科化石在四川的首次发现．     

广西石炭兽科新材料

本文记述了产自广西南宁盆地中部长岭坡(村)竹排冲岸边的石炭兽一新种:长岭石炭兽(Anthracotherium changlingensis sp. nov.)这是我国目前发现的体型最大的石炭兽。通过对化石的记述和对比,证明我国有Anthracotherium这一属的存在,并确定了南宁盆地邕宁组上段(即上含煤段)的地质时代为晚渐新世。文中还记述了百色盆地发现的两件石炭兽牙齿化石。     

THE DYNAMICS OF WOUND CLOSURE AND ITS ROLE IN THE PROGRAMMING OF PLANARIAN REGENERATION. II — DISTALIZATION

When the dorsal and ventral epidermal layers join by first intention during the closure of the wound, the cells of their borders (M-cells) do not meet in the same manner in all sections. In anterior sections the dorsal M-cells attach themselves to the ventral basement membrane, so that only the dorsal epidermis is stretched. In posterior sections the dorsal and the ventral M-cells join by their apical edges without being closely apposed to the wound surface. Only the ventral cells are stretched because of their specific motility. In longitudinal sections the dorsal and the ventral M-cells also join by their apical edges, but since they are closely apposed to the wound surface both epidermal layers are stretched. The stretching is a process equivalent to distalization. The junction between the dorsal and the ventral epidermis is shifted ventrally in the anterior wounds (as in the intact heads) and dorsally in the posterior wounds (as in the intact tails). Some abnormalities of wound closure have been observed at levels where heteromorphic regeneration frequently occurs. These findings are consistent with the hypothesis previously advanced (3) that the modalities of wound closure establish the programme for regeneration.     

COMPARATIVE ULTRASTRUCTURAL STUDIES OF SPERMATOGENESIS IN THE METZGERIALES (HEPATOPHYTA) II. THE BLEPHAROPLAST OF BLASIA PUSILLA

As in other hepatics, the young spermatid of Blasia pusilla contains a well-developed blepharoplast comprising a four-layered multilayered structure (MLS) and two overlying dimorphic basal bodies. The asymmetrical spline (S₁ or upper stratum of the MLS) numbers 20 or 21 microtubules (MTs) at its anterior tip and reduces to eight at the posterior limit of the lamellar strip (LS). Behind this the shank of the spline is five or six tubules in width over most of its length, approximately one revolution of the circumference of the gamete. The three-microtubule spline aperture underlies the anterior basal body and like those of most hepatics, it is closed at its anterior end. The asymmetrical LS (approx. 2.0 μm in length) is characterized by a right-hand posterior notch which lies below the spline aperture at the region of the cartwheel configuration of the anterior basal body (ABB). The staggered dimorphic basal bodies overlap for approximately one third of their lengths. Both lie parallel to the long axis of the spline. As in other hepatics, the ABB (1.2 μm in length) is subapical and comprises an anterior hub extension with progressive rearward additions of lateral, dorsal and ventral triplets. Over most of its length (2.1 μm) the longer posterior basal body (PBB) consists of a distinct central hub and three ventral triplets. Transition zones of both basal bodies contain stellate configurations into which the two central axonemal MTs frequently extend. The blepharoplast of Blasia shows several features in common with leafy, simple thalloid and complex thalloid liverworts. Compared with the few Metzgeriales observed thus far, the LS is less elongate and the basal bodies less staggered. Dimensions of basal body components and spline dimensions, however, are comparable to those of most leafy and thalloid hepatics. Striking similarities with the complex thalloid liverworts include a posterior notch in the LS and a spline aperture three MTs wide.     

THE MICROTUBULAR CYTOSKELETON OF AMPHIDINIUM RHYNCHOCEPHALUM (DINOPHYCEAE)1

The sub-thecal microtubular cytoskeleton of Amphidinium rhynchocephalum Anissimowa was investigated using indirect immunofluorescence microscopy and transmission electron microscopy. The majority of sub-thecal microtubules are longitudinally oriented and radiate from one of two sub-thecal transverse microtubular bands that lie adjacent to the anterior and posterior edge of the cingulum.Both transverse bands consist of 3–5 microtubules and are loop shaped with one end adjacent to the cell's right edge of the sulcus and the other end adjacent to the fibrous ventral ridge. The posterior transverse microtubular band (PTB) defines the posterior edge of the cingulum and gives rise to numerous posteriorly directed longitudinal microtubular bundles that consist of 1–3 microtubules per bundle. These bundles end at the posterior end of the cell. The PTB also gives rise to the cingular longitudinal microtubules that underlie the cingular groove and terminate at the anterior transverse microtubular band (ATB). The ATB defines the anterior edge of the cingulum and loops around the base of the epicone. This band gives rise to anteriorly directed longitudinal microtubular bundles that terminate in the small epicone of the cell. The longitudinal microtubular root of the flagellar apparatus is directed posteriorly and lies immediately beneath the theca but is distinct from the subthecal microtubule system. A narrow fibrous ridge is ventrally located to the cell's left between the exit apertures of the transverse and longitudinal flagella. In this position, the ventral ridge lies between and also connects with the anterior and posterior transverse microtubular bands. The ventral ridge is also associated with three microtubules that are distinct from other cytoskeletal microtubules. Our results demonstrate that the majority of sub-thecal microtubules originate from one of two microtubular bands associated with the cingulum. The possible role of the fibrous ventral ridge and its associated microtubules is also discussed.     

固定化微藻对虾池弧菌数量动态的影响

引入固定化波吉卵囊藻(Oocystis borgei)和微绿球藻(Nannochloris oculata)于凡纳对虾(Litop Penaeus vannamei)养殖环境中,检测水体、对虾胃和后肠中弧菌的数量变化,研究固定化微藻对虾池弧数量动态影响。结果表明:波吉卵囊藻培养液中9d后不能检测出弧菌,微绿球藻培养液中15d后不能检测出弧菌。引入固定化波吉卵囊藻和微绿球藻的褐藻胶藻珠能抑制弧菌的生长,实验组养殖水体、对虾胃和后肠中弧菌的数量都比对照组低;抑制效果是固定化波吉卵囊藻和微绿球藻混合固定化波吉卵囊藻固定化微绿球藻;试验后期实验组弧菌的数量明显低于试验前期。试验期间固定化波吉卵囊藻和微绿球藻的生物量分别增加了约10倍和17倍,证明它们的生理活性不会因固定化而受干扰。因此,固定化微藻可应用于虾池微生态调控防病。