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1.
The morphological organization and functional anatomy of prosomal defensive (scent) glands in Paranemastoma quadripunctatum, a representative of the dyspnoid harvestmen, was investigated by means of histological semithin sections, software‐based 3D‐reconstruction and scanning electron microscopy. Scent glands comprise large, hollow sacs on either side of the prosoma, each of these opening to the outside via one orifice (ozopore) immediately above coxa I. In contrast to the situation known from laniatorean, cyphophthalmid and some eupnoid Opiliones, ozopores are not exposed but hidden in a depression (atrium), formed by a dorsal integumental fold of the carapace and the dorsal parts of coxae I. Glandular sacs are connected to ozopores via a short duct which is equipped with a specific closing mechanism in its distal part: A layer of modified epidermal cells forms a kind of pad‐like tissue, surrounding the duct like a valve. Several muscles attached to the anterior parts of the glandular reservoir and to the epithelial pad may be associated with ozopore‐opening. The actual mechanism of secretion discharge seems to be highly unusual and may be hypothesized on the basis of corroborating data from behavioral observations, scent gland anatomy and secretion chemistry as follows: Enteric fluid is considered to be directed towards the ozopores via cuticular grooves in the surface of the coxapophyses of legs I. Then, the fluid is sucked into the anterior part of the scent gland reservoirs by the action of dorsal dilator muscles that widen the reservoir and produce a short‐term negative pressure. After dilution/solution of the naphthoquinone‐rich scent gland contents, a secretion‐loaded fluid is thought to be discharged with the help of transversal compressor muscles. This is the first detailed study on the functional anatomy of scent glands and the mechanisms of secretion discharge in the Dyspnoi. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.  相似文献   

2.
Females of a solitary digger wasp, the European beewolf (Philanthus triangulum F.), cultivate symbiotic bacteria of the genus Streptomyces in specialized antennal glands. The streptomycetes are secreted in the subterranean brood cells and protect the offspring against mould fungi. We reconstructed the complex morphology of the antennal glands using 3D-visualization software, investigated the ultrastructure of the glands, and examine the role of the antennal glands as organs for the cultivation of the symbiotic bacteria. The bacteria are cultivated in five antennomeres within large reservoirs that consist of two slightly bent lobes. Each gland reservoir is bordered by a monolayered epithelium lined with a partially reinforced cuticle and when completely filled with bacteria it comprises about half of the antennomere's volume. The opening of the reservoir is covered by gelatinous appendage of the cuticle. The cells of the monolayered epithelium bordering each reservoir show basal invaginations, apical microvilli and numerous vesicles. Each reservoir is surrounded by approximately 400 class 3 gland units that are connected to the reservoir lumen through conducting canals. The class 3 gland cells contain numerous vesicles and a high density of rough endoplasmatic reticulum. In the reservoir lumen, large numbers of symbiotic Streptomyces bacteria are embedded in secretion droplets. Thus, the bacteria are apparently provided with large amounts of nutrients via the gland epithelium and the class 3 gland cell units.  相似文献   

3.
Movement of fluids through geological porous media is an important factor in ore genesis, crude oil genesis, and in oil recovery from oil reservoirs. Such a movement may be beneficially affected by biological in situ activity. We have shown that once bacteria have been introduced into a porous medium, they are able to penetrate into their environment and multiply without decreasing significantly the rock's permeability. The limiting bacterial concentration, which may cause plugging, depends on the bacterial size, pore size, and pore size distribution of the reservoir rock. Microorganisms may, under anaerobic conditions, affect the composition of crude oil by increasing its asphaltenic fraction. In simulated systems microorganisms increased the recovery of the oil in place approximately five times. A bacterial consortium was more effective than a single pure bacterial strain.  相似文献   

4.
Summary The Middle-Upper Jurassic section in the Arabian Gulf basin forms one of the most prolific sequences in the world, in which an excellent combination of source, reservoir and seal rocks was developed within a major sedimentary cycle. The sequence consists of a) relatively quiet deep-water mudstone, wackestone and shale (source facies), b) shallow-water high enery grainstone and packstone (reservoir facies), and c) very shallow supratidal anhydrite (seal facies). The principal factors, which controlled the sedimentation of this sequence, are considered to have been eustatic sea-level change and epeirogenic movement of carbonate shelves. The Jurassic reservoirs of the major oil fields in this region show exceptionally high porosity up to 30% for their relatively old geologic age (some 150 million years old) and depths of burial in the range between 1,200 and more than 2,700 m. Porosity occurs most commonly as intergranular/remnant primary pore spaces, but its distribution is quite uneven and very complicated. To account for the existence of such high porosity (and permeability) in the Jurassic reservoirs, probable geological, physical and chemical factors for preserving and enhancing porosity (and permeability), such as acidic formation fluids, reduced fluid mobility, tectonic forces, ductility of intercalated beds (e.g. anhydrite), and dolomitization were examined. It has been observed in various fields in the region that oilsaturated portions of the Jurassic reservoirs tend to retain higher porosity than the surrounding water-saturated zones. Porosity preservation by hydrocarbons is possible primarily because of excess hydrocarbon pressure and of reduced mobility of water in such oil-saturated zones. To continue sediment diagenesis, a steady supply of minerals by formation water and the mobility of the water may have been essential. Because the entrapment of oil in the Jurassic reservoirs in the region is considered to have been as late as early Tertiary, some other (pre-migration) mechanisms which may have worked in the earlier geologic stages for preserving and creating porosity (and permeability) seem to be necessary.  相似文献   

5.
The sternum V glands are a key synapomorphy that unites Trichoptera with Lepidoptera, but their functional aspects have not been analysed from an evolutionary perspective. We examine phylogenetic trends and correlations between chemical and morphological features of these glands. The most likely ancestral gland compounds are heptan‐2‐ol, 4‐hepten‐2‐one and ‐ol, nonan‐2‐one, and 6‐nonen‐2‐one and ‐ol, making pheromone production a plausible ancestral function. The most widespread gland compounds (heptan‐2‐one and ‐ol and nonan‐2‐one and ‐ol) are not known from Apataniidae + Limnephilidae (Trichoptera), which in turn uniquely produce a number of methylated 3‐ketones and their corresponding alcohols, probably functioning as pheromones. We propose a functional connection between perforated patches on sternum IV in females and a scaly/dome‐covered area around the gland openings, as well as between perforated patches and lack of Trichoptera‐type opening muscles. We also propose a functional connection between the shape of the gland reservoirs and the presence of gland reservoir musculature. The perforated patches were significantly correlated with several gland compounds that had double bonds between carbon atoms: the double bonds may lower the viscosity of the compounds, facilitating secretion through the tiny pores of the perforated patches. The production of defensive substances in Pycnopsyche (Trichoptera: Limnephilidae) is probably connected to the presence of large, compartmentalized gland reservoirs. Large glands in male Hydropsyche (Trichoptera: Hydropsychidae) are probably linked to male aggregation pheromone production. The relative sizes of sternum V gland reservoirs and fenestral gland reservoirs in female philopotamids (Trichoptera) suggest a complementary function of the two structures. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 561–579.  相似文献   

6.
7.
目前我国油田开发主要处于高含水后期,微生物驱提高石油采收率技术(MEOR)以低成本、环境友好等独特的优势引起了石油工业界的重视。实际上,经过半个多世纪的发展,MEOR已经成为提高采收率的重要前沿技术。高压是油藏的主要环境特征,在影响油藏微生物生存与活性等方面具有重要作用。本文从油藏及其微生物的主要特征、微生物对高压环境的适应机制以及高压下微生物降解烃的代谢特征等方面进行了综述。介绍了对油藏微生物资源、群落结构、微生物在油水相中分布的认识,微生物乳化原油机制,以及微生物在油藏厌氧环境中协同代谢、受温度和压力影响的特点,并列举了MEOR的矿场应用。在高压适应机制上,微生物主要通过改变和调整细胞膜结构、增加胞内脂质组分和表达胞内特殊酶等作用来实现对压力的适应;在高压下烃降解微生物代谢速率低于常压,而且耐压菌和嗜压菌具有不同的烃降解效率。  相似文献   

8.
Biological souring and mitigation in oil reservoirs   总被引:1,自引:0,他引:1  
Souring in oilfield systems is most commonly due to the action of sulfate-reducing prokaryotes, a diverse group of anaerobic microorganisms that respire sulfate and produce sulfide (the key souring agent) while oxidizing diverse electron donors. Such biological sulfide production is a detrimental, widespread phenomenon in the petroleum industry, occurring within oil reservoirs or in topside processing facilities, under low- and high-temperature conditions, and in onshore or offshore operations. Sulfate reducers can exist either indigenously in deep subsurface reservoirs or can be “inoculated” into a reservoir system during oilfield development (e.g., via drilling operations) or during the oil production phase. In the latter, souring most commonly occurs during water flooding, a secondary recovery strategy wherein water is injected to re-pressurize the reservoir and sweep the oil towards production wells to extend the production life of an oilfield. The water source and type of production operation can provide multiple components such as sulfate, labile carbon sources, and sulfate-reducing communities that influence whether oilfield souring occurs. Souring can be controlled by biocides, which can non-specifically suppress microbial populations, and by the addition of nitrate (and/or nitrite) that directly impacts the sulfate-reducing population by numerous competitive or inhibitory mechanisms. In this review, we report on the diversity of sulfate reducers associated with oil reservoirs, approaches for determining their presence and effects, the factors that control souring, and the approaches (along with the current understanding of their underlying mechanisms) that may be used to successfully mitigate souring in low-temperature and high-temperature oilfield operations.  相似文献   

9.
About half of the world's oil production is from carbonate formations. However, most of the research in microbially enhanced oil recovery (MEOR), a potentially important tertiary recovery technology, has focused on sandstone reservoirs because, in general, they are geologically simpler than carbonate reservoirs and easier to model in the laboratory. Carbonate formations have a wide range of pore geometries and distributions, resulting in complex flow dynamics. The low matrix permeabilities and the dual porosity characteristics of most carbonate formations, coupled with the chemistry of carbonates, have slowed implementation of enhanced oil recovery methods. A review of the data on carbonate reservoirs in Dwight's Energydata TOTL System indicated that 40% of the oil‐producing carbonate reservoirs surveyed in the United States have environmental, geological, and petrophysical conditions that would make them candidates for MEOR. A review of a number of MEOR field trials showed that rates of oil production could be increased by as much as 200%. Microbial activity in these trials was probably due to that of indigenous populations rather than the microorganisms injected for the trials. Detrimental effects such as loss of injectivity and increased souring were not reported. Based on analysis of the geology and petrophysical characteristics of carbonates, two common mechanisms of MEOR, microbial acid production and microbial gas production, are especially suited for application in carbonate reservoirs.  相似文献   

10.
Oil reservoirs represent a nutrient-rich ecological niche of the deep biosphere. Although most oil reservoirs are occupied by microbial populations, when and how the microbes colonized these environments remains unanswered. To address this question, we compared 11 genomes of Thermotoga maritima-like hyperthermophilic bacteria from two environment types: subsurface oil reservoirs in the North Sea and Japan, and marine sites located in the Kuril Islands, Italy and the Azores. We complemented our genomes with Thermotoga DNA from publicly available subsurface metagenomes from North America and Australia. Our analysis revealed complex non-bifurcating evolutionary history of the isolates'' genomes, suggesting high amounts of gene flow across all sampled locations, a conjecture supported by numerous recombination events. Genomes from the same type of environment tend to be more similar, and have exchanged more genes with each other than with geographically close isolates from different types of environments. Hence, Thermotoga populations of oil reservoirs do not appear isolated, a requirement of the ‘burial and isolation'' hypothesis, under which reservoir bacteria are descendants of the isolated communities buried with sediments that over time became oil reservoirs. Instead, our analysis supports a more complex view, where bacteria from subsurface and marine populations have been continuously migrating into the oil reservoirs and influencing their genetic composition. The Thermotoga spp. in the oil reservoirs in the North Sea and Japan probably entered the reservoirs shortly after they were formed. An Australian oil reservoir, on the other hand, was likely colonized very recently, perhaps during human reservoir development.  相似文献   

11.
Bacterial communities in both aqueous and oil phases of water-flooded petroleum reservoirs were characterized by molecular analysis of bacterial 16S rRNA genes obtained from Shengli Oil Field using DNA pyrosequencing and gene clone library approaches. Metagenomic DNA was extracted from the aqueous and oil phases and subjected to polymerase chain reaction amplification with primers targeting the bacterial 16S rRNA genes. The analysis by these two methods showed that there was a large difference in bacterial diversity between the aqueous and oil phases of the reservoir fluids, especially in the reservoirs with lower water cut. At a high phylogenetic level, the predominant bacteria detected by these two approaches were identical. However, pyrosequencing allowed the detection of more rare bacterial species than the clone library method. Statistical analysis showed that the diversity of the bacterial community of the aqueous phase was lower than that of the oil phase. Phylogenetic analysis indicated that the vast majority of sequences detected in the water phase were from members of the genus Arcobacter within the Epsilonproteobacteria, which is capable of degrading the intermediates of hydrocarbon degradation such as acetate. The oil phase of reservoir fluid samples was dominated by members of the genus Pseudomonas within the Gammaproteobacteria and the genus Sphingomonas within the Alphaproteobacteria, which have the ability to degrade crude oil through adherence to hydrocarbons under aerobic conditions. In addition, many anaerobes that could degrade the component of crude oil were also found in the oil phase of reservoir fluids, mainly in the reservoir with lower water cut. These were represented by Desulfovibrio spp., Thermodesulfovibrio spp., Thermodesulforhabdus spp., Thermotoga spp., and Thermoanaerobacterium spp. This research suggested that simultaneous analysis of DNA extracted from both aqueous and oil phases can facilitate a better understanding of the bacterial communities in water-flooded petroleum reservoirs.  相似文献   

12.
Prothoracic and abdominal defensive glands are present in the genus Menimus (Coleoptera; Tenebrionidae). The prothoracic glands are small and open laterally into the membrane between the prothorax and the head. The reservoirs of the abdominal glands exceed the length of the abdomen and the secretory tubules end separately in ‘basal lines’ dorsally on the base of each reservoir. Chemical analysis of the glandular secretions revealed the presence of alkylated benzoquinones, ethylphenol and undecene. The relationships of the genus Menimus are discussed.  相似文献   

13.
Gas transport in unconventional shale strata is a multi-mechanism-coupling process that is different from the process observed in conventional reservoirs. In micro fractures which are inborn or induced by hydraulic stimulation, viscous flow dominates. And gas surface diffusion and gas desorption should be further considered in organic nano pores. Also, the Klinkenberg effect should be considered when dealing with the gas transport problem. In addition, following two factors can play significant roles under certain circumstances but have not received enough attention in previous models. During pressure depletion, gas viscosity will change with Knudsen number; and pore radius will increase when the adsorption gas desorbs from the pore wall. In this paper, a comprehensive mathematical model that incorporates all known mechanisms for simulating gas flow in shale strata is presented. The objective of this study was to provide a more accurate reservoir model for simulation based on the flow mechanisms in the pore scale and formation geometry. Complex mechanisms, including viscous flow, Knudsen diffusion, slip flow, and desorption, are optionally integrated into different continua in the model. Sensitivity analysis was conducted to evaluate the effect of different mechanisms on the gas production. The results showed that adsorption and gas viscosity change will have a great impact on gas production. Ignoring one of following scenarios, such as adsorption, gas permeability change, gas viscosity change, or pore radius change, will underestimate gas production.  相似文献   

14.
Ptychoidy is a defensive adaptation of several groups of oribatid mites in which legs and coxisternum can be fully retracted into the opisthosoma and protected by a ventrally deflected prodorsum, resulting in a seed-like appearance. Using Euphthiracarus cooki as a model, we examined details of exoskeletal and muscular anatomy in combination with studies of live individuals to provide the first functional analysis of ptychoidy. There are two main functional components: the first is a set of exoskeletal and muscular adaptations, mostly of the podosoma and prodorsum, that combine to effect leg withdrawal and prodorsal deflection; the second comprises adaptations of the opisthosoma that allow control of hydrostatic pressure during the large hemocoel volume adjustments associated with ptychoidy. Adaptations important in the closing process (enptychosis) are found in four body regions. Much of the podosomal exoskeleton (especially pleural) is unsclerotized, which facilitates leg retraction and prodorsal deflection during enptychosis. The coxisternum has several flexible furrows along which it folds in order to bring legs into a tightly parallel arrangement. The prodorsum has specialized attachment surfaces (manubrium and inferior retractor process) for retractor muscles and a paired bothridial scale that participates in prodorsal alignment during enptychosis. The subcapitulum has a prominent capitular apodeme on which important retractor muscles insert. The mineralized notogaster has an anterior "collar" that accommodates the retracted prodorsum; it includes paired notches and receptacles that accommodate the bothridial scales, thereby creating a temporary fixed axis for rotation of the prodorsum in a "lazy hinge" mechanism. Specialized muscles form the retractor system; most conspicuous are the large coxisternal retractors and prodorsal retractors, both of which originate on the notogaster. Other components have adjustor roles; among them are muscles of the endosternal system that control retraction of the subcapitulum and assist leg retraction, and the dorsoventral muscles which adjust the folded coxisternum. Hemolymph pressure control is a function of the opisthosoma, where the principle exoskeletal elements form a pleated venter, having a cross-sectional shape like an inverted "W." Paired holoventral plates (each representing fused genital, aggenital, anal, and anal plates) form the inner angle and are flanked by paired plicature plates. The holoventral plates are connected medially in two ways : 1) by three permanent bridges of sclerotized cuticle that include an anterior phragmatal bridge and two widely spaced, hollow apodemes (preanal, postanal); 2) by temporary zipper-like closures of two different types. Lateral compression of the pleats is effected by a series of transversely arranged, lateral compressor muscles that run from plicature to holoventral plate edges, and from holoventral plate edges to the medial apodemes. Compression increases hydrostatic pressure in the opisthosoma and stores energy in both the slightly deformed, mineralized notogaster and in the three holoventral bridges. During normal activity the compressor system is active and the inflated podosomal region provides support for the extended legs. When the mite is irritated, the prodorsum is hydraulically ejected from its active position in the notogastral collar, then relaxation of the compressors causes a fall in hemolymph pressure and return of the notogaster to an undeformed condition. Muscles of the retractor system then act in specific sequence to retract and adjust the coxisternum and prodorsum until they are precisely positioned at the completion of enptychosis. The process takes between 0.5 and 1 sec. When irritation ceases, partial opening allows sensory leg hairs to "test" the environment. Resumption of normal, active posture (ecptychosis) involves activation of the lateral compressor system and hydraulic inflation of the podosoma, through which legs are extended and the prodorsum is reflected.  相似文献   

15.
Stimulated reservoir volume (SRV) fracturing in tight oil reservoirs often induces complex fracture-network growth, which has a fundamentally different formation mechanism from traditional planar bi-winged fracturing. To reveal the mechanism of fracture network propagation, this paper employs a modified displacement discontinuity method (DDM), mechanical mechanism analysis and initiation and propagation criteria for the theoretical model of fracture network propagation and its derivation. A reasonable solution of the theoretical model for a tight oil reservoir is obtained and verified by a numerical discrete method. Through theoretical calculation and computer programming, the variation rules of formation stress fields, hydraulic fracture propagation patterns (FPP) and branch fracture propagation angles and pressures are analyzed. The results show that during the process of fracture propagation, the initial orientation of the principal stress deflects, and the stress fields at the fracture tips change dramatically in the region surrounding the fracture. Whether the ideal fracture network can be produced depends on the geological conditions and on the engineering treatments. This study has both theoretical significance and practical application value by contributing to a better understanding of fracture network propagation mechanisms in unconventional oil/gas reservoirs and to the improvement of the science and design efficiency of reservoir fracturing.  相似文献   

16.
Microbial enhanced oil recovery (MEOR) refers to the process of using bacterial activities for more oil recovery from oil reservoirs mainly by interfacial tension reduction and wettability alteration mechanisms. Investigating the impact of these two mechanisms on enhanced oil recovery during MEOR process is the main objective of this work. Different analytical methods such as oil spreading and surface activity measurements were utilized to screen the biosurfactant-producing bacteria isolated from the brine of a specific oil reservoir located in the southwest of Iran. The isolates identified by 16S rDNA and biochemical analysis as Enterobacter cloacae (Persian Type Culture Collection (PTCC) 1798) and Enterobacter hormaechei (PTCC 1799) produce 1.53 g/l of biosurfactant. The produced biosurfactant caused substantial surface tension reduction of the growth medium and interfacial tension reduction between oil and brine to 31 and 3.2 mN/m from the original value of 72 and 29 mN/m, respectively. A novel set of core flooding tests, including in situ and ex situ scenarios, was designed to explore the potential of the isolated consortium as an agent for MEOR process. Besides, the individual effects of wettability alteration and IFT reduction on oil recovery efficiency by this process were investigated. The results show that the wettability alteration of the reservoir rock toward neutrally wet condition in the course of the adsorption of bacteria cells and biofilm formation are the dominant mechanisms on the improvement of oil recovery efficiency.  相似文献   

17.

Laboratory evaluation of hyperthermophiles with the potential for Enhanced Oil Recovery (EOR) is often hampered by the difficulties in replicating the in situ growth conditions in the laboratory. In the present investigation, genome analysis was used to gain insights into the metabolic potential of a hyperthermophile to mobilize the residual oil from depleting high-temperature oil reservoirs. Here, we report the 1.9 Mb draft genome sequence of a hyperthermophilic anaerobic archaeon, Thermococcus sp. 101C5, with a GC content of 44%, isolated from a high-temperature oil reservoir of Gujarat, India. 101C5 possessed the genetic arsenal required for adaptation to harsh oil reservoir conditions, such as various heat shock proteins for thermo-adaptation, Trk potassium uptake system proteins for osmo-adaptation, and superoxide reductases against oxidative stress. Microbial Enhanced Oil Recovery (MEOR) potential of the strain was established by ascertaining the presence of genes encoding enzymes involved in the production of the metabolites such as hydrogen, bio-emulsifier, acetate, exopolysaccharide, etc. Production of these metabolites which pressurize the reservoir, emulsify the crude oil, lower the viscosity and reduce the drag, thus facilitating mobilization of the residual oil was experimentally confirmed. Also, the presence of crude oil degradative genes highlighted the ability of the strain to mobilize heavy residual oil, which was confirmed under simulated conditions in sand-pack studies. The obtained results demonstrated additional oil recoveries of 42.1% and 56.5% at 96 °C and 101 °C, respectively, by the strain 101C5, illustrating its potential for application in high-temperature oil reservoirs. To our best knowledge, this is the first report of genome analysis of any microbe assessed for its suitability for MEOR from the high-temperature oil reservoir.

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18.
19.
The construction of large reservoirs in South America and particularly in Brazil has intensified within the last 50 years. Built up primarily for hydroelectricity production, these artificial ecosystems now serve purposes such as: water storage for public use; fisheries and aquaculture; recreation; tourism, and irrigation. These artificial ecosystems were also built up with the purpose of enhancing the regional development. These activities produce multiple impacts among which are eutrophication, a serious problems with various ecological, economic, and social consequences. Basic studies on reservoirs have identified their main ecological characteristics and described some of their fundamental mechanisms of functioning. This information cover: factors involved in reservoir complexity, such as spatial scale, vertical and horizontal heterogeneity, temporal variation at several time scales and, in some cases, watershed/reservoir relationships, and the impact of non-point and point sources of nutrients. Numerous studies have been dedicated to describing aquatic biota, its relationship with environmental factors, and the impacts of degradation/pollution/ eutrophication on biodiversity. Research priorities have been established mainly by limnologists in response to needs identified by environmental impact assessment. Due to the pressure of multiple uses, population growth, and economic factors, aquatic scientists now face a number of questions being posed by managers and engineers. These questions relate to a wide range of practical problems, the solutions to which depend on accumulated data related to the structural complexities and function mechanisms previously referred to. These problems can be divided into three main groups: (i) eutrophication processes and their characteristics, and water quality control; (ii) impact of fisheries, aquaculture, and exotic species introduction on the biota and the water quality; (iii) sustainable development of reservoirs and optimization management of their multiple uses. Predictive capabilities required in the face of these problems should be based on modeling development and intensive use of data-bases resulting from long-term studies on reservoirs. The predictions thus made possible would involve economic factors and the costs of recovery of eutrophic reservoirs. The experience of reservoir research and management in South America and in Brazil is, probably part of a world wide tendency on reservoir research and development with the aim to optimize multiple uses. Based on the cooperation of limnologists and engineers, ecosystem management models must eventually be applied at the watershed / reservoir level, where demands on the part of managers and decision makers will certainly promote a predictive, integrated, and adaptive approach,based on fundamental research.  相似文献   

20.
The oesophagus of Nereis virens is divided into anterior and posterior regions and possesses several peculiar features not previously described for cuticle or glands of Polychaetes either in body or foregut integument. The mucous cells which are permanent glands in both regions secrete a carboxylated acidic mucus and possess apical mechanisms permitting the opening and closing of the neck of the glands. The cuticle of the posterior oesophagus bears numerous striated spines and several hypotheses concerning their physiological role are discussed.  相似文献   

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