共查询到20条相似文献,搜索用时 19 毫秒
1.
R T Kimball E L Braun J D Ligon 《Proceedings. Biological sciences / The Royal Society》1997,264(1387):1517-1523
Afropavo congensis, the Congo peafowl, has long fascinated ornithologists because of its uncertain phylogenetic position and unusual geographic distribution. While some researchers have placed Afropavo as a sister taxon to the true peafowl, Pavo species, others have suggested relationships with the guineafowl or an Old World partridge, Francolinus. These divergent opinions are due, at least in part, to (i) the unique morphological characteristics, lack of elaborate ornamentation, and monogamous mating system in Afropavo which differentiates it from Pavo; and (ii) the restricted distribution of Afropavo in Zaire, which is far removed from the Asian distribution of all other pheasant species. We obtained complete cytochrome b and partial D-loop sequences of Afropavo and compared them to Pavo, guineafowl, Francolinus and other galliform taxa. Our results strongly support a close relationship between Afropavo and Pavo, and we were able to reject alternative phylogenetic hypotheses. Molecular clock estimates of the divergence time place the separation of Afropavo and Pavo in the late Miocene. We also discuss other relatives of Afropavo and Pavo and use this information to propose hypotheses regarding the evolution of ornamentation and sexual dimorphism within this group of pheasants. 相似文献
2.
Petrov NB Pegova AN Manylov OG Vladychenskaia NS Miuge NS Aleshin VV 《Molekuliarnaia biologiia》2007,41(3):499-507
Gastrotrichs are meiobenthic free-living aquatic worms whose phylogenetic and intra-group relationships remain unclear despite some attempts to resolve them on the base of morphology or molecules. In this study we analysed complete sequences of the 18S rRNA gene of 15 taxa (8 new and 7 published) to test numerous hypotheses on gastrotrich phylogeny and to verify whether controversial interrelationships from previous molecular data could be due to the short region available for analysis and the poor taxa sampling. Data were analysed using both maximum likelihood and Bayesian inference. Results obtained suggest that gastrotrichs, together with Gnathostomulida, Plathelminthes, Syndermata (Rotifera + Acanthocephala), Nemertea and Lophotrochozoa, comprise a clade Spiralia. Statistical tests reject phylogenetic hypotheses regarding Gastrotricha as close relatives of Nematoda and other Ecdysozoa or placing them at the base of bilaterian tree close to acoels and nemertodermatides. Within Gastrotricha, Chaetonotida and Macrodasyida comprise two well supported clades. Our analysis confirmed the monophyly of the Chaetonotidae and Xenotrichulidae within Chaetonida as well as Turbanellidae and Thaumastodermatidae within Macrodasyida. Mesodasys is a sister group of the Turbanellidae, and Lepidodasyidae appears to be a polyphyletic group as Cephalodasys forms a separate lineage at the base of macrodasyids, whereas Lepidodasys groups with Neodasys between Thaumastodermatidae and Turbanellidae. To infer a more reliable Gastrotricha phylogeny many species and additional genes should be involved in future analyses. 相似文献
3.
Despite the wide range of shapes and sizes that accompany a vast variety of functions, the development of tetrapod limbs follows a conservative pattern of de novo condensation, branching, and segmentation. Development of the zeugopodium and digital arch typically occurs in a posterior to anterior sequence, referred to as postaxial dominance, with a digital sequence of 4-3-5-2-1. The only exception to this pattern in all of living Tetrapoda can be found in salamanders, which display a preaxial dominance in limb development, a de novo condensation of a basale commune (distal carpal/tarsal 1+2) and a precoccial development of digits I and II. These divergent patterns have puzzled researchers for over a century leading to various explanatory hypotheses. Despite many advances in research on tetrapod limb development, the divergent evolution of these two pathways and its causes are still not understood. Based on an extensive ontogenetic series we investigated the pattern of limb development of the 300 Ma old branchiosaurid amphibian Apateon. This revealed a preaxial dominance in limb development that was previously believed to be unique and derived for modern salamanders. The Branchiosauridae are favored as close relatives of extant salamanders in most phylogenetic hypotheses of the highly controversial origins and relationships of extant amphibians. The findings provide new insights into the evolution of developmental pathways in tetrapod limb development, the relationships of modern amphibians with possible Paleozoic antecedents, and their initial timing of divergence. 相似文献
4.
Fuchs J Fjeldså J Bowie RC Voelker G Pasquet E 《Molecular phylogenetics and evolution》2006,39(1):186-197
The African genus Hyliota includes three or four species of warbler-like birds of uncertain phylogenetic affinities, as it has historically been placed in different avian families that are now known to represent unrelated lineages: Malaconotidae (bush-shrikes), Platysteiridae (batises and wattle-eyes), Muscicapidae (Old World flycatchers) and Sylviidae (Old-World Warblers). To assess the affinities of Hyliota we sequenced a mitochondrial protein-coding gene (ND2, 1018bp) and a nuclear intron (myoglobin intron-2, 685bp). Our analyses suggest that all previous hypotheses concerning the affinities of Hyliota are erroneous. Instead, Hyliota represents a basal branch in the Passerida radiation with no close relatives. Our results, which also include analyses of relationships among other of other atypical songbird genera, lend support to an African origin of the Passerida songbird radiation. 相似文献
5.
Recent hypotheses argue that phylogenetic relatedness should predict both the niche differences that stabilise coexistence and the average fitness differences that drive competitive dominance. These still largely untested predictions complicate Darwin's hypothesis that more closely related species less easily coexist, and challenge the use of community phylogenetic patterns to infer competition. We field parameterised models of competitor dynamics with pairs of 18 California annual plant species, and then related species' niche and fitness differences to their phylogenetic distance. Stabilising niche differences were unrelated to phylogenetic distance, while species' average fitness showed phylogenetic structure. This meant that more distant relatives had greater competitive asymmetry, which should favour the coexistence of close relatives. Nonetheless, coexistence proved unrelated to phylogeny, due in part to increasing variance in fitness differences with phylogenetic distance, a previously overlooked property of such relationships. Together, these findings question the expectation that distant relatives should more readily coexist. 相似文献
6.
The objectives of this study were to: (1) describe diffusion of information by affected women in whom a mutation has been identified (index cases) to their families and testing participation among high-risk relatives; (2) assess information recall and understanding by index cases and their satisfaction with the testing process; and (3) determine the factors associated with higher/lower testing decision in the family. Thirty index cases completed a self-administered questionnaire assessing their personal and family characteristics and their satisfaction with their own genetic testing process and a telephone interview to evaluate their knowledge about the risk of a genetic predisposition to breast and ovarian cancer, the type and number of close relatives that they informed, and the difficulties that they encountered. Information about breast/ovarian cancer risk and test availability was generally well transmitted (75%), predominantly (88%) to first-degree relatives. In contrast, testing participation was low (15%) and essentially occurred among sisters and daughters. There was a general lack of knowledge despite a high level of satisfaction regarding the information given by the geneticist. Family support and the knowledge of index cases about the risk of transmission of BRCA1/2 mutations by women were found to be positively and significantly associated with the testing decision among first-degree relatives. Difficulties in informing relatives appeared to be related to poor understanding of the information by index cases, as well as fear, and avoidance among close relatives. A major challenge for genetic counseling is to ensure that consulting patients not only receive complete understanding but also understand this information and anticipate the impact of the test result before deciding to take the test. 相似文献
7.
Mainland colonization by island lizards 总被引:4,自引:0,他引:4
Kirsten E. Nicholson Richard E. Glor † Jason J. Kolbe Allan Larson S. Blair Hedges Jonathan B. Losos 《Journal of Biogeography》2005,32(6):929-938
Aim We investigate biogeographic relationships within the lizard genus Anolis Daudin, 1802 to test the hypothesis that the mainland (Central and South American) Norops‐clade species descended from a West Indian Anolis ancestor. Previous hypotheses have suggested that close island relatives of mainland Norops species (the Cuban Anolis sagrei and Jamaican A. grahami series) represent over‐water dispersal from a mainland ancestor. These previous hypotheses predict that the A. sagrei and A. grahami series should be phylogenetically nested within a Norops clade whose ancestral geography traces to the mainland. If Norops is West Indian in origin, then West Indian species should span the deepest phylogenetic divergences within the Norops clade. Location Central and South America and West Indian islands. Methods The phylogenetic relationships of Anolis lizards are reconstructed from aligned DNA sequences using both parsimony and Bayesian approaches. Hypotheses are tested in two ways: (1) by reconstructing the ancestral geographic location for the Norops clade using Pagel & Lutzoni's (2002) Bayesian approach, and (2) by testing alternative topological arrangements via Wilcoxon Signed‐Ranks tests ( Templeton, 1983 ) and Shimodaira–Hasegawa tests ( Shimodaira & Hasegawa, 1999 ). Results Our evidence supports an origin of mainland Norops anoles from a West Indian ancestor. A West Indian ancestor to the Norops clade is statistically supported, and alternatives to the biogeographic pattern [Cuban (Jamaican, Mainland)] are statistically rejected by Shimodaira–Hasegawa tests, although not by Wilcoxon Signed‐Ranks tests. Main conclusions Our data support the hypothesis of a West Indian origin for mainland Norops. This result contradicts previous hypotheses and suggests that island forms may be an important source for mainland biodiversity. 相似文献
8.
C. Kik A. M. Samoylov W. H. J. Verbeek L. W. D. van Raamsdonk 《TAG. Theoretical and applied genetics. Theoretische und angewandte Genetik》1997,94(3-4):465-471
Mitochondrial (mt) DNA variation in the cultigens leek, kurrat and prei-anak is limited compared to that of their wild relatives
in the Allium ampeloprasum complex. The phylogenetic relationships among these cultigens and their wild relatives is quite close, with the majority
of the species clustering within one mitochondrial clade. The presence in leek of an extra-mitochondrial genetic element was
noted. Analysis of crossability showed that all species were interfertile with leek. It is suggested that the genetic variation
present within the A. ampeloprasum complex could be exploited in order to broaden the genetic basis of leek.
Received: 14 August 1996 / Accepted: 23 August 1996 相似文献
9.
Predicting invasion potential has global significance for managing ecosystems as well as important theoretical implications for understanding community assembly. Phylogenetic relationships of introduced species to the extant community may be predictive of establishment success because of the opposing forces of competition/shared enemies (which should limit invasions by close relatives) versus environmental filtering (which should allow invasions by close relatives). We examine here the association between establishment success of introduced birds and their phylogenetic relatedness to the extant avifauna within three highly invaded regions (Florida, New Zealand, and Hawaii). Published information on both successful and failed introductions, as well as native species, was compiled for all three regions. We created a phylogeny for each avifauna including all native and introduced bird species. From the estimated branch lengths on these phylogenies, we calculated multiple measurements of relatedness between each introduced species and the extant avifauna. We used generalized linear models to test for an association between relatedness and establishment success. We found that close relatedness to the extant avifauna was significantly associated with increased establishment success for exotic birds both at the regional (Florida, Hawaii, New Zealand) and sub‐regional (islands within Hawaii) levels. Our results suggest that habitat filtering may be more important than interspecific competition in avian communities assembled under high rates of anthropogenic species introductions. This work also supports the utility of community phylogenetic methods in the study of vertebrate invasions. 相似文献
10.
Efe Pygmies in northeast Zaire exhibit features of social organization (small group size, flexible but predominantly patrilocal residence, close relations with kin, and pair bonds) that are common to many hunting and gathering societies. This study uses methods commonly employed in studies of nonhuman primates to investigate the associations of Efe men with other individuals as a function of their age, sex, and kinship, and it tests the hypothesis that Efe are patrilocal because hunting efficiency is improved by hunting with male relatives. The analyses of 376 hours of focal behavior observations on 16 Efe men and observations of 71 cooperative group hunts show that the majority of associations between Efe men were with other adult men. Men associated preferentially with kin over nonkin, and they associated with close kin more than with distant kin. Men's close relationships (what we call “Companionships” sensu Smuts' “Friendships” in baboons) were predominantly with other adult men, but each man who cohabited with a woman had his strongest Companionship by far with that woman. Neither the efficiency of hunts nor the hunting success of individual men were found to be related to the degree of relatedness of hunters nor the proportion of relatives on the hunt. Alternative hypotheses concerning the functional significance of patrilocality and male kin groups are considered. We conclude that strong affiliative bonds between male kin and between males and females may arise as much from the need for strong allies in the face of competitive social situations as from economic or ecological necessity. 相似文献
11.
The hypothesis that outcrossers are competitively superior to close relatives with specialized genetic systems was tested. Interspecific competition between three closely related Oenothera species was examined in common garden environments. Oenothera grandis, an obligate outcrosser, is competitively superior to its derivative, O. laciniata, an obligate chromosomal heterozygote. Both of these species are superior to O. mexicana, a closely related facultative selfer. These relationships were consistent over two levels of density stress. Therefore, in this species triad, the outcrosser has a competitive advantage over its close relatives with derived breeding systems. 相似文献
12.
Mitochondrial DNA sequence data from the control region and 12S rRNA in leopard frogs from the Sierra El Aguaje of southern Sonora, Mexico, together with GenBank sequences, were used to infer taxonomic identity and provide phylogenetic hypotheses for relationships with other members of the Rana pipiens complex. We show that frogs from the Sierra El Aguaje belong to the Rana berlandieri subgroup, or Scurrilirana clade, of the R. pipiens group, and are most closely related to Rana magnaocularis from Nayarit, Mexico. We also provide further evidence that Rana magnaocularis and R. yavapaiensis are close relatives. 相似文献
13.
The evolution of migration in birds has fascinated biologists for centuries. In this study, we performed phylogenetic-based analyses of Catharus thrushes, a model genus in the study of avian migration, and their close relatives. For these analyses, we used both mitochondrial and nuclear genes, and the resulting phylogenies were used to trace migratory traits and biogeographic patterns. Our results provide the first robust assessment of relationships within Catharus and relatives and indicate that both mitochondrial and autosomal genes contribute to overall support of the phylogeny. Measures of phylogenetic informativeness indicated that mitochondrial genes provided more signal within Catharus than did nuclear genes, whereas nuclear loci provided more signal for relationships between Catharus and close relatives than did mitochondrial genes. Insertion and deletion events also contributed important support across the phylogeny. Across all taxa included in the study, and for Catharus, possession of long-distance migration is reconstructed as the ancestral condition, and a North American (north of Mexico) ancestral area is inferred. Within Catharus, sedentary behaviour evolved after the first speciation event in the genus and is geographically and temporally correlated with Central American distributions and the final closure of the Central American Seaway. Migratory behaviour subsequently evolved twice in Catharus and is geographically and temporally correlated with a recolonization of North America in the late Pleistocene. By temporally linking speciation events with changes in migratory condition and events in Earth history, we are able to show support for several competing hypotheses relating to the geographic origin of migration. 相似文献
14.
Evolutionary changes in color patches of blackbirds are associated with marsh nesting 总被引:2,自引:0,他引:2
Fully unraveling the mechanisms of sexual selection requiresan understanding of the variation in secondary sexual traitsacross species in a monophyletic assemblage and an understandingof the evolutionary relationships between those species. Therole of red and yellow male plumage coloration in territorydefense and sexual selection has been well studied in the red-winged
blackbird (Agelaius phoeniceus), and males of many other close
relatives of this species also have what appear to be carotenoid-pigmented
patches in their plumage. We explored variation in male plumagecoloration across species of New World blackbirds (family Icteridae):traits known to be involved in sexual selection in this group.We document that blackbird lineages in which extant speciesbreed in marshes tend to have evolved from an all-black ancestralplumage to one exhibiting carotenoid plumage patches. The twomost likely hypotheses to explain this pattern are (1) increasedsexual selection intensity in marshes because of increasedvariance in territory quality and (2) increased frequency ofmale-male territorial interactions because of an increaseddensity of territories in marshes, but other hypotheses cannotbe ruled out. This pattern is consistent with either intersexualor intrasexual selection and warrants further investigation. 相似文献
15.
Aim Geckos (Reptilia: Squamata), due to their great age and global distribution, are excellent candidates to test hypotheses of Gondwanan vicariance against post‐Gondwanan dispersal. Our aims are: to generate a phylogeny of the sphaerodactyl geckos and their closest relatives; evaluate previous phylogenetic hypotheses of the sphaerodactyl geckos with regard to the other major gecko lineages; and to use divergence date estimates to inform a biogeographical scenario regarding Gondwanan relationships and assess the roles of vicariance and dispersal in shaping the current distributions of the New World sphaerodactyl geckos and their closest Old World relatives. Location Africa, Asia, Europe, South America, Atlantic Ocean. Methods We used parsimony and partitioned Bayesian methods to analyse data from five nuclear genes to generate a phylogeny for the New World sphaerodactyl geckos and their close Old World relatives. We used dispersal–vicariance analysis to determine ancestral area relationships among clades, and divergence times were estimated from the phylogeny using nonparametric rate smoothing. Results We recovered a monophyletic group containing the New World sphaerodactyl genera, Coleodactylus, Gonatodes, Lepidoblepharis, Pseudogonatodes and Sphaerodactylus, and the Old World Gekkotan genera Aristelliger, Euleptes, Quedenfeldtia, Pristurus, Saurodactylus and Teratoscincus. The dispersal–vicariance analysis indicated that the ancestral area for this clade was North Africa and surrounding regions. The divergence between the New World spaherodactyl geckos and their closest Old World relative was estimated to have occurred c. 96 Myr bp . Main conclusions Here we provide the first molecular genetic phylogenetic hypothesis of the New World sphaerodactyl geckos and their closest Old World relatives. A combination of divergence date estimates and dispersal–vicariance analysis informed a biogeographical scenario indicating that the split between the sphaerodactyl geckos and their African relatives coincided with the Africa/South America split and the opening of the Atlantic Ocean. We resurrect the family name Sphaerodactylidae to represent the expanded sphaerodactyl clade. 相似文献
16.
Rauscher JT 《American journal of botany》2002,89(7):1074-1084
The subtribe Espeletiinae (Asteraceae, Heliantheae) comprises morphologically and ecologically diverse plants endemic to the tropical montane paramos of the Andes of Venezuela, Colombia, and Ecuador. Though the ecophysiology and ecology of this adaptive radiation have been well studied, relationships among taxa in the subtribe and between the subtribe and other taxa in the Heliantheae are poorly known. In this study, sequences from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA are used to test previous hypotheses about the phylogenetic position of the Espeletiinae within the Heliantheae and to determine which taxa are the subtribe's closest relatives. Gene phylogenies based on maximum parsimony analyses reveal that the Espeletiinae clade is nested well within the subtribe Melampodiinae and thus should be considered a monophyletic complex of species, not a separate subtribe. The most parsimonious gene trees suggest that the genus Ichthyothere may be the sister taxon to the Espeletia complex and that the genus Smallanthus and a species of Rumfordia are likely among the complex's other closest living relatives. These data offer preliminary insights into the origins of this adaptive radiation and the broader phylogenetic context in which it occurred. 相似文献
17.
The low body mass index (BMI) phenotype of less than 18.5 has been linked to medical and psychological morbidity as well as increased mortality risk. Although genetic factors have been shown to influence BMI across the entire BMI, the contribution of genetic factors to the low BMI phenotype is unclear. We hypothesized genetic factors would contribute to risk of a low BMI phenotype. To test this hypothesis, we conducted a genealogy data analysis using height and weight measurements from driver''s license data from the Utah Population Data Base. The Genealogical Index of Familiality (GIF) test and relative risk in relatives were used to examine evidence for excess relatedness among individuals with the low BMI phenotype. The overall GIF test for excess relatedness in the low BMI phenotype showed a significant excess over expected (GIF 4.47 for all cases versus 4.10 for controls, overall empirical p-value<0.001). The significant excess relatedness was still observed when close relationships were ignored, supporting a specific genetic contribution rather than only a family environmental effect. This study supports a specific genetic contribution in the risk for the low BMI phenotype. Better understanding of the genetic contribution to low BMI holds promise for weight regulation and potentially for novel strategies in the treatment of leanness and obesity. 相似文献
18.
An extremely large number of fifth ceratobranchial teeth, with highly modified, striated, and hooked tips were observed in the central and western Pacific sicydiine goby genus Stiphodon.A scanning electron microscopic study of the form and arrangement of fifth ceratobranchial teeth was conducted to assess the distribution of these modifications in sicydiine gobies and their putative close relatives. Our goals were to explore a new set of characters in gobioid systematics, to test sicydiine monophyly, and to test hypotheses of relationships of sicydiine gobies. Sicydiines are hypothesized herein to be most closely related to the western Pacific Tukugobius and Rhinogobius,freshwater genera with which they share thickened pelvic-fin rays, no teeth on the anterior portion of the fifth ceratobranchial bones, fifth ceratobranchial teeth with differentiated and striated tips, and overlapping anterior rami of the fifth ceratobranchial bones. The latter two characters occur in some, but not all, sicydiines. The pantropical freshwater goby Awaous,often classified with sicydiines, is not considered the closest relative of the subfamily. The highly modified fifth ceratobranchials of Stiphodon are similar to, and concluded here to be homoplasious with, those of the mudflat-dwelling New World goby Evorthodus and the Indo-west Pacific oxudercine gobies, represented in this study by Pseudapocryptes. J. Morphol. 237:257–274, 1998. Published 1998 Wiley-Liss, Inc. 相似文献
19.
Morphological and palynological studies in Phlomis L., Eremostachys Bunge and Paraphlomis Prain (Labiatae) 总被引:1,自引:0,他引:1
DINA AZIZIAN D. M. MOORE F.L.S. 《Botanical journal of the Linnean Society. Linnean Society of London》1982,85(4):225-248
Information from morphology and palynology, together with anatomical, cytological and chemical data, is utilized to examine the status of the genus Phlomis and its close relatives Eremostachys and Paraphlomis.
The recognition of the genus Paraphlomis is confirmed on morphological and palynological grounds. The data also reinforce earlier opinions about the close relationship between Phlomis and Eremostachys, which show some overlap in almost all the features examined. When infrageneric groups are considered, the indumentum, calyx-shape and pollen-sculpturing in Eremostachys section Moluccelloides differ markedly from these features in Eremostachys sections Eremostachys and Metaxoides, which show close affinities with Phlomis, especially with Phlomis section Phlomoides. Phlomis sections Phlomis and Phlomoides differ in morphology, petiolar anatomy, chromosome number and constituent flavonols. Palynological data, allied to those from morphology and geography, strongly support the removal of the Chinese species, Phlomis rolata, to the genus Lamiophlomis. 相似文献
The recognition of the genus Paraphlomis is confirmed on morphological and palynological grounds. The data also reinforce earlier opinions about the close relationship between Phlomis and Eremostachys, which show some overlap in almost all the features examined. When infrageneric groups are considered, the indumentum, calyx-shape and pollen-sculpturing in Eremostachys section Moluccelloides differ markedly from these features in Eremostachys sections Eremostachys and Metaxoides, which show close affinities with Phlomis, especially with Phlomis section Phlomoides. Phlomis sections Phlomis and Phlomoides differ in morphology, petiolar anatomy, chromosome number and constituent flavonols. Palynological data, allied to those from morphology and geography, strongly support the removal of the Chinese species, Phlomis rolata, to the genus Lamiophlomis. 相似文献
20.
Rudall PJ Remizowa MV Prenner G Prychid CJ Tuckett RE Sokoloff DD 《American journal of botany》2009,96(1):67-82
Reproductive units (RUs) of Trithuria, the sole genus of the early-divergent angiosperm family Hydatellaceae, are compared with flowers of their close relatives in Cabombaceae (Nymphaeales). Trithuria RUs combine features of flowers and inflorescences. They differ from typical flowers in possessing an "inside-out" morphology, with carpels surrounding stamens; furthermore, carpels develop centrifugally, in contrast to centripetal or simultaneous development in typical flowers. Trithuria RUs could be interpreted as pseudanthia of two or more cymose partial inflorescences enclosed within an involucre, but the bractlike involucral phyllomes do not subtend partial inflorescences and hence collectively resemble a typical perianth. Teratological forms of T. submersa indicate a tendency to fasciation and demonstrate that the inside-out structure-the primary feature that separates RUs of Hydatellaceae from more orthodox angiosperm flowers-can be at least partially modified, thus producing a morphology that is closer to an orthodox flower. The Trithuria RU could be described as a "nonflower", i.e., a structure that contains typical angiosperm carpels and stamens but does not allow recognition of a typical angiosperm flower. The term nonflower could combine cases of secondary loss of flower identity and cases of a prefloral condition, similar to those that gave rise to the angiosperm flower. Nonhomology among some angiosperm flowers could be due to iterative shifts between nonfloral construction and flower/inflorescence organization of reproductive organs. Potential testing of these hypotheses using evolutionary-developmental genetics is explored using preliminary data from immunolocalization of the floral meristem identity gene LEAFY in T. submersa, which indicated protein expression at different hierarchical levels. 相似文献