106 resultados para Less favoured regions


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Morphologies of solution-cast films of iPP/aPP blends have been studied by means of electron microscopy and X-ray scattering techniques. Microscopic observation showed that solution-cast film of iPP consists of two kinds of structural regions, cross-hatched and lath-liked structures. The addition of small amount of aPP (less than or equal to 30%) into iPP did not change iPP's characteristic crystallization behavior. It is noticed that when the content of aPP in its blend was over 80%, iPP formed a very loosely woven-like network composed of very long lamellae with wide-angle lamellar branchings. The X-ray data showed that aPP did not cocrystallize with iPP.

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The unicellular green alga Haematococcus pluvialis accumulates a commercially valuable astaxanthin, with levels reaching up to 4% dry weight under environmental stress. In recent years, much effort has been devoted to understanding the molecular mechanisms regulating astaxanthin biosynthetic pathways. Beta-carotene ketolase (bkt), with control being exhibited at the transcription level, plays an important role in astaxanthin biosynthesis by H. pluvialis. Here we demonstrate the presence of two separate 5'-flanking regions [1.5 kilobase (kb) and 2 kb] of bkt (bkt1 and bkt2) that possess regulatory elements similar to those of known stress-responsive genes in plants. Results of 5'-deletion constructs and transient beta-galactosidase expression assays demonstrate that there may be positive regulatory elements governing expression in the shorter promoter at -1060/-900 from the 1.5 kb 5' region, and in the longer promoter at -1838/-1219 and at -1046/ -734 from the 2 kb 5' region relative to each homologous ATG start codon. Furthermore, our present studies reveal that the first intron (+371/+497) downstream from the 1.5 kb 5' untranslated region of bkt1 may function as a negative regulatory element to regulate its own promoter.

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The complete mitochondrial (mt) genome sequence of Oratosquilla oratoria (Crustacea: Malacostraca: Stomatopoda) was determined; a circular molecule of 15,783 bp in length. The gene content and arrangement are consistent with the pancrustacean ground pattern. The mt control region of O. oratoria is characterized by no GA-block near the 3' end and different position of [TA(A)]n-blocks compared with other reported Stomatopoda species. The sequence of the second hairpin structure is relative conserved which suggests this region may be a synapomorphic character for the Stomatopoda. In addition, a relative large intergenic spacer (101 bp) with higher A + T content than that in control region was identified between the tRNA(Glu) and tRNA(Phe) genes. Phylogenetic analyses based on the current dataset of complete mt genomes strongly support the Stomatopoda is closely related to Euphausiacea. They in turn cluster with Penaeoidea and Caridea clades while other decapods form a separate group, which rejects the monophyly of Decapoda. This challenges the suitability of Stomatopoda as an outgroup of Decapoda in phylogenetic analyses. The basal position of Stomatopoda within Eumalacostraca according to the morphological characters is also questioned. (C) 2010 Elsevier Inc. All rights reserved.

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This study investigates the distribution of black carbon (BC) and its correlation with total polycyclic aromatic hydrocarbons (I PAH) pound in the surface sediments of China's marginal seas. BC content ranges from < 0.10 to 2.45 mg/g dw (grams dry weight) in the sediments studied, and varied among the different coastal regions. The Bohai Bay sediments had the highest BC contents (average 2.18 mg/g dw), which comprises a significant fraction (27%-41%) of the total organic carbon (TOC) preserved in the sediments. In comparison, BC in the surface sediments of the North Yellow Sea, Jiaozhou Bay, East China Sea and the South China Sea is less abundant and accounted for an average of 6%, 8%, 14% and 5%, respectively, of the sedimentary organic carbon pool. The concentration of I PAH pound in the surface sediments ranges from 41 to 3 667 ng/g dw and showed large spatial variations among the sampling sites of different costal regions. The Bohai Bay has the highest I PAH pound values, ranging from 79 to 3 667 ng/g dw. This reflects the high anthropogenically contaminated nature of the sediments in the bay. BC is positively correlated to TOC but a strong correlation is not found between BC and I PAH pound in the surface sediments studied, suggesting that BC and PAHs preserved in the sediments are derived from different sources and controlled by different biogeochemical processes. Our study suggests that the abundance of BC preserved in the sediments could represent a significant sink pool of carbon cycling in China's marginal seas.

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We used fifteen years (1993-2007) of altimetric data, combined from different missions (ERS-1/2, TOPEX/Poseidon, Jason-1, and Envisat), to analyze the variability of the eddy kinetic energy (EKE) in the South China Sea (SCS). We found that the EKE ranged from 64 cm(2)/s(2) to 1 390 cm(2)/s(2) with a mean value of 314 cm(2)/s(2). The highest EKE center was observed to the east of Vietnam (with a mean value of 509 cm(2)/s(2)) and the second highest EKE region was located to the southwest of Taiwan Island (with a mean value of 319 cm(2)/s(2)). We also found that the EKE structure is the consequence of the superposition of different variability components. First, interannual variability is important in the SCS. Spectral analysis of the EKE interannual signal (IA-EKE) shows that the main periodicities of the IA-EKE to the east of Vietnam, to the southwest of Taiwan Island, and in the SCS are 3.75, 1.87, and 3.75 years, respectively. It is to the south of Taiwan Island that the IA-EKE signal has the most obvious impact on EKE variability. In addition, the IA-EKE exhibit different trends in different regions. An obvious positive trend is observed along the east coast of Vietnam, while a negative trend is found to the southwest of Taiwan Island and in the east basin of Vietnam. Correlation analysis shows that the IA-EKE has an obvious negative correlation with the SSTA in Nio3 (5A degrees S-5A degrees N, 90A degrees W-150A degrees W). El Nio-Southern Oscillation (ENSO) affects the IA-EKE variability in the SCS through an atmospheric bridge-wind stress curl over the SCS. Second, the seasonal cycle is the most obvious timescale affecting EKE variability. The locations of the most remarkable EKE seasonal variabilities in the SCS are to the east of Vietnam, to the southwest of Taiwan, and to the west of Philippines. To the east of Vietnam, the seasonal cycle is the dominant mechanism controlling EKE variability, which is attributed primarily to the annual cycle there of wind stress curl. In this area, the maximum EKE is observed in autumn. To the southwest of Taiwan Island, the EKE is enlarged by the stronger SCS circulation, which is caused by the intrusion branch from the Kuroshio in winter. Finally, intra-annual and mesoscale variability, although less important than the former, cannot be neglected. The most obvious intra-annual and mesoscale variability, which may be the result of baroclinic instability of the background flow, are observed to the southwest of Taiwan Island. Sporadic events can have an important effect on EKE variability.

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Based on the data analysis, this study further explores the characteristics of East Asian winter monsoon (hereafter, EAWM, for brevity) as well as the related air-sea-land system, and illustrates how and to what degree anomalous signals of the subsequent Asian summer monsoon are rooted in the preceding EAWM activity. We identified an important air-sea coupled mode, i.e., the EAWM mode illustrated in Section 3. In cold seasons, strong EAWM-related air-sea two-way interaction is responsible for the development and persistence of the SSTA pattern of EAWM mode. As a consequence, the key regions, i.e., the western Pacific and South China Sea (hereafter, SCS, for brevity), are dominated by such an SSTA pattern from the winter to the following summer. In the strong EAWM years, the deficient snow cover dominates eastern Tibetan Plateau in winter, and in spring, this anomaly pattern is further strengthened and extended to the northwestern side of Tibetan Plateau. Thus, the combined effect of strong EAWM-related SSTA and Tibetan snow cover constitutes an important factor in modulating the Asian monsoon circulation. The active role of the EAWM activity as well as the related air-sea-land interaction would, in the subsequent seasons, lead to: 1) the enhancement of SCS monsoon and related stronger rainfall; 2) the northward displacement of subtropical high during Meiyu period and the related deficient rainfall over Meiyu rainband; 3) above-normal precipitation over the regions from northern Japan to northeastern China in summer; 4) more rainfall over the Arabian Sea and Northeast India, while less rainfall over southwest India and the Bay of Bengal. The strong EAWM-related air-sea interaction shows, to some degree, precursory signals to the following Asian summer monsoon. However, the mechanism for the variability of Indian summer monsoon subsequent to the strong EAWM years remains uncertain.

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利用ERA40逐日再分析资料、NCEP/NCAR2逐日再分析资料、中国740个测站日降水资料、上海台风研究所提供的西太平洋热带气旋资料、Kaplan等重建的月平均SSTA资料、NOAA逐日长波辐射(OLR)等资料,应用离散功率谱分析、带通滤波、EOF分析等统计方法,研究了东亚夏季风(EASM)的移动特征、东亚地区季节内振荡(ISO)的基本特征、季节内振荡对东亚夏季风活动的影响、季节内振荡对东亚夏季风异常活动的影响机理。主要结论如下: (1)综合动力和热力因素定义了可动态描述东亚夏季风移动和强度的指数,并利用该指数研究了东亚夏季风的爆发和移动的季节内变化及其年际和年代际变化特征。研究发现,气候平均东亚夏季风前沿分别在28候、33候、36候、38候、40候、44候出现了明显的跳跃。东亚夏季风活动具有显著的年际变率,主要由于季风前沿在某些区域异常停滞和突然跨越北跳或南撤引起,造成中国东部旱涝灾害频繁发生。东亚夏季风的活动具有明显的年代际变化,在1965年、1980年、1994年发生了突变,造成中国东部降水由“南旱北涝”向“南涝北旱”的转变。 (2)东亚季风区季节内变化具有10~25d和30~60d两个波段的季节内振荡周期,以30-60d为主。存在三个主要低频模态,第一模态主要表征了EASM在长江中下游和华北地区活动期间的低频形势;第二模态印度洋-菲律宾由低频气旋式环流控制,主要表现了ISO在EASM爆发期间的低频形势;第三模态主要出现在EASM在华南和淮河活动期间的低频形势。第一模态和第三模态是代表东亚夏季风活动异常的主要低频形势。 (3)热带和副热带地区ISO总是沿垂直切变风的垂直方向传播。因此,在南海-菲律宾东北风垂直切变和副热带西太平洋北风垂直切变下,大气热源激发菲律宾附近交替出现的低频气旋和低频反气旋不断向西北传播,副热带西太平洋ISO以向西传播为主。中高纬度地区,乌拉尔山附近ISO以向东、向南移动或局地振荡为主;北太平洋中部ISO在某些情况下向南、向西传播。 (4)季风爆发期,伴随着热带东印度洋到菲律宾一系列低频气旋和低频反气旋, 冷空气向南输送,10~25天和30~60天季节内振荡低频气旋同时传入南海加快了南海夏季风的爆发。在气候态下,ISO活动表现的欧亚- 太平洋(EAP)以及太平洋-北美(PNA)低频波列分布特征(本文提出的EAP和PNA低频波列与传统意义上的二维定点相关得到的波列不同)。这种低频分布形式使得欧亚和太平洋中高纬度的槽、脊及太平洋副热带高压稳定、加强,东亚地区的低频波列则成为热带和中高纬度ISO相互作用影响东亚夏季风活动的纽带。不同的阶段表现不同的低频模态,30~60d低频模态的转变加快了EASM推进过程中跳跃性;30-60d低频模态的维持使得EASM前沿相对停滞。 (5)30-60d滤波场,菲律宾海域交替出现的低频气旋和低频反气旋不断向西北传播到南海-西太平洋一带。当南海-西太平洋地区低频气旋活跃时,季风槽加强、东伸,季风槽内热带气旋(TC)频数增加;当南海-西太平洋低频反气旋活跃时,季风槽减弱、西退,TC处于间歇期,生成位置不集中。 (6)在El Nino态下,大气季节内振荡偏弱,北传特征不明显,但ISO由中高纬度北太平洋中部向南和副热带西太平洋向西的传播特征显著,东亚地区ISO活动以第三模态为主,EASM集中停滞在华南和淮河流域,常伴随着持续性区域暴雨的出现,易造成华南和江淮流域洪涝灾害,长江和华北持续干旱。在La Nina态下,大气季节内振荡活跃,且具有明显的向北传播特征,PNA低频波列显著,东亚地区ISO活动以第一模态单峰为主;EASM主要停滞在长江中下游和华北地区,这些地区出现异常持续强降水,华南和淮河流域多干旱;在El Nino态向La Nina态转换期,ISO活动以第一模态双峰为主,长江中下游常常出现二度梅。

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The morphology and infraciliature of two ectoparasitic ciliates, Trichodina caecellae n. sp. and T. ruditapicis Xu, Song & Warren, 2000, parasitising the gills of marine molluscs from the Shandong coast of the Yellow Sea, China, were investigated following wet silver nitrate and protargol impregnation. T. caecellae was found on the small marine sand clam Caecella chinensis Deshayes and is distinguished mainly by the acute triangle-like blade, the very delicate central part and the needle-shaped ray. T. ruditapicis was studied based on four populations from three clams: two populations from Ruditapes philippinarum (Adams) and one each from Saxidomus purpuratus (Sowerby) and Solen grandis Dunker. All four populations fell within the range of morphometry and agreed closely in the overall appearance of the adhesive disc. However, variability was found in the denticle structure, especially in populations from different host clams. Our observations suggest that denticle morphology may be more or less variable between and within populations, and that such minor differences should not be overestimated. It should be emphasised that, except for the denticle morphology, the bright granules or circles in the centre of the adhesive disc represent another important feature facilitating the identification of this trichodinid species.

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The complete mitochondrial (mt) DNA sequence was determined for a ridgetail white prawn, Exopalaemon carinicauda Holthuis, 1950 (Crustacea: Decopoda: Palaemonidae). The mt genome is 15,730 bp in length, encoding a standard set of 13 protein-coding genes, 2 ribosomal RNA genes, and 22 transfer RNA genes, which is typical for metazoans. The majority-strand consists of 33.6% A, 23.0% C, 13.4% G, and 30.0% T bases (AT skew = 0.057: GC skew = -0.264). A total of 1045 bp of non-coding nucleotides were observed in 16 intergenic regions,,including a major A+ T rich (79.7%) noncoding region (886 bp). A novel translocation of tRNA(Pro) and tRNA(Thr) was found when comparing this genome with the pancrustacean ground pattern indicating that gene order is not conserved among caridean mitochondria. Furthermore, the rate of Ka/Ks in 13 protein-coding genes between three caridean species is Much less than 1, which indicates a strong Purifying selection within this group. To investigate the phylogenetic relationship within Malacostraca, phylogenetic trees based oil Currently available malacostracan complete mitochondrial sequences were built with the maximum likelihood and Bayesian models. All analyses based oil nucleotide and amino acid data strongly support the monophyly of Decapoda. The Penaeidae, Reptantia, Caridea, and Meiura clades were also recovered as monophyletic groups with Strong Statistical Support. However, the phylogenetic relationships within Pleocyemata are unstable, as represented by the inclusion or exclusion of Caridea. (C) 2009 Elsevier B.V. All rights reserved.

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Geographic and vertical variations of size-fractionated (0.2-1 mu m, 1-10 mu m, and >10 mu m) Chlorophyll a (Chl.a) concentration, cyanobacteria abundance and heterotrophic bacteria abundance were investigated at 13 stations from 4 degrees S, 160 degrees W to 30 degrees N, 140 degrees E in November 1993. The results indicated a geographic distribution pattern of these parameters with instances of high values occurring in the equatorial region and offshore areas, and with instance of low values occurring in the oligotrophic regions where nutrients were almost undetectable. Cyanobacteria showed the highest geographic variation (ranging from 27x10(3) to 16,582x10(3) cell l(-1)), followed by Chl.a (ranging from 0.048 to 0.178 mu g l(-1)), and heterotrophic bacteria (ranging from 2.84x10(3) to 6.50 x 10(5) cell l(-1)). Positive correlations were observed between nutrients and Chl.a abundance. Correspondences of cyanobacteria and heterotrophic bacteria abundances to nutrients were less significant than that of Chl.a. The total Chl.a was accounted for 1.0-30.9%, 35.9-53.7%, and 28.1-57.3% by the >10 mu m, 1-10 mu m and 0.2-1 mu m fractions respectively. Correlation between size-fractionated Chl.a and nutrients suggest that the larger the cell size, the more nutrient-dependent growth and production of the organism. The ratio of pheophytin to chlorophyll implys that more than half of the > 10 mu m and about one third of the 1-10 mu m pigment-containing particles in the oligotrophic region were non-living fragments, while most of the 1-10 mu m fraction was living cells. In the depth profiles, cyanobacteria were distributed mainly in the surface layer, whereas heterotrophic bacteria were abundant from surface to below the euphotic zone. Chl.a peaked at the surface layer (0-20 m) in the equatorial area and at the nitracline (75-100 m) in the oligotrophic regions. Cyanobacteria were not the principle component of the picoplankton. The carbon biomass ratio of heterotroph to phytoplankton was greater than 1 in the eutrophic area and lower than 1 in oligotrophic waters.

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Polyploidization plays an important role in generating the current high diversity of plants. Studies of distributional patterns of diploids and derivative autopolyploids have provided important insights into evolutionary processes and cryptic speciation of polyploidization within species defined on the basis of their morphology. However, few studies have been designed to examine distributions of infrageneric diploids and polyploids on the Asian Qinghai-Tibetan Plateau (QTP). Allium przewalskianum occurs widely on the QTP and in adjacent regions, at altitudes ranging from 2000m to 4500m. We collected a total of 844 individuals from 62 populations and determined their cytotypes over the entire distribution range of this species. Tetraploids tend to occur at high altitudes; however, the positive relationship between the ploidy and altitude was only marginally significant (P < 0.05). Contact zones between diploids and tetraploids were recorded on the eastern QTP from north to south. Four populations were found to harbor both cytotypes, but no triploid individuals. The wider distribution of tetraploids may be mainly due to their greater colonization ability in the new niches created by the Quaternary climatic oscillations in the QTP region. Our results offer a fundamental framework for studying evolutionary origins, adaptations and cryptic divergences of polyploids within this species complex based on molecular and/or ecological examinations in the future.

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Background and Aims It is an enduring question as to the mechanisms leading to the high diversity and the processes producing endemics with unusual morphologies in the Himalayan alpine region. In the present study, the phylogenetic relationships and origins of three such endemic genera were analysed, Dolomiaea, Diplazoptilon and Xanthopappus, all in the tribe Cardueae of Asteraceae.Methods The nuclear rDNA internal transcribed spacer (ITS) and plastid trnL-F and psbA-trnH regions of these three genera were sequenced. The same regions for other related genera in Cardueae were also sequenced or downloaded from GenBank. Phylogenetic trees were constructed from individual and combined data sets of the three types of sequences using maximum parsimony, maximum likelihood and Bayesian analyses.Key Results The phylogenetic tree obtained allowed earlier hypotheses concerning the relationships of these three endemic genera based on gross morphology to be rejected. Frolovia and Saussurea costus were deeply nested within Dolomiaea, and the strong statistical support for the Dolomiaea-Frolovia clade suggested that circumscription of Dolomiaea should be more broadly redefined. Diplazoptilon was resolved as sister to Himalaiella, and these two together are sister to Lipschitziella. The clade comprising these three genera is sister to Jurinea, and together these four genera are sister to the Dolomiaea-Frolovia clade. Xanthopappus, previously hypothesized to be closely related to Carduus, was found to be nested within a well-supported but not fully resolved Onopordum group with Alfredia, Ancathia, Lamyropappus, Olgaea, Synurus and Syreitschikovia, rather than the Cardinis group. The crude dating based on ITS sequence divergence revealed that the divergence time of Dolomiaea-Frolovia from its sister group probably occurred 13.6-12.2 million years ago (Ma), and the divergence times of the other two genera, Xanthopappus and Diplazoptilon, from their close relatives around 5.7-4.7 Ma and 2.0-1.6 Ma, respectively.Conclusions The findings provide an improved understanding of the intergeneric relationships in Cardueae. The crude calibration of lineages indicates that the uplifts of the Qiinghai -Tibetan Plateau since the Miocene might have served as a continuous stimulus for the production of these morphologically aberrant endemic elements of the Himalayan flora.

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Heavy mineral assemblages, chemical compositions of diagnostic heavy minerals such as garnet and tourmaline, and U-Pb ages and Hf isotopic compositions of zircons are very effective means to determine sediment provenance. An integrated application of the above provides insight on the lithologies, crystallization ages and crustal formation ages of the parent magma of sediment source areas. As a result, the locations and characteristics of potential source areas can be constrained and contributions of different source regions may be evaluated. In addition, the study provides evidence for the magmatic and tectonic history of source areas using a novel approach. The heavy mineral assemblages, and chemical compositions of detrital garnets and tourmalines, U-Pb ages and Hf isotopic compositions of zircons for sand and loess samples deposited since the Last Glacial Maximum (LGM) from the Hulunbeier, Keerqin and Hunshandake sandlands were analyzed and compared to those of central-southern Mongolia, the central Tarim and surrounding potential source areas, the Central Asian Orogenic Belt (CAOB) and North China Craton (NCC). The following remarks on provenance and tectonic history can be made: 1. The source compositional characteristics of the Hulunbeier, Keerqin and Hunshandake sandlands are similar. They are derived from the CAOB and NCC whose contributions for the Keerqin and Hunshandake sandland are about 50%. For the Hulunbeier sandland it is somewhat less, about 40%. 2. Loesses around of the sandlands have the identical source signiture as the sands, implying that they are sorted by the same wind regime. 3. The source characteristics of the present and LGM sands are the same, providing direct evidence that the present sands originated from the reworking of LGM sands. 4. The provenance characteristics of the three sandlands differ from those of the Tarim. As a result, the possibility that the three eastern sandlands were sourced from the Taklimakan desert can be ruled out. 5. The source compositions of sand samples derived from the CAOB indicate that the occurrence of Archean and Paleoproterozoic metamorphic basement rocks is limited and continuous subduction-accretion events from the Neoproterozoic to the Mesozoic occurred. This implies that the CAOB is a orogenic collage belt similar to the present day southwest-Pacific, and formed by the amalgamation of small forearc and backarc ocean basins occurring between island arcs and microcontinents during continuous collision and accretion. The Hf isotopic signitures of detrital zircons indicate that large amounts of juvenile mantle materials were added to the CAOB crust during the Phanerozoic.

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The West Shandong Uplift and its adjacent basins, with same evolutional history before Mesozoic, are an important basin-orogenic systems in North China. After late Mesozoic, tectonic differentiation between basin and orogenic belt gradually displayed in the study area. The Boxing sag is a part of Jiyang Depression near to West Shandong Uplift, in which the whole Mesozoic and Cenozoic strata are preserved. Based on the analysis of sedimentary records in the Boxing sag, the Cenozoic structural and sedimentary evolutions in Boxing Sag and its response to Western Shandong uplift are discussed in this dissertation. The main conclusions in this research are presented as follows. Based on Seismic and well logging profile interpretation, fault growth index, thickness difference between bottom wall and top wall and fault activity rate from Eocene to Pliocene are studied. Boxing sag had three main faults, NE, NW and NEE trending faults. Research shows that the activity of the NW trending fault in the Boxing sag became weaken from E1-2S4 to N2m gradually. The evolution of NE and the NEE trending fault can be divided into three episodes, from E1-2k to E2s4, from E2s3 to E3s1, from N2m to E3d. The analysis of Paleogene samples of heavy mineral assemblages shows that metamorphic rocks represented by garnet, intermediate-acid igneous rocks represented by the assemblage of apatite, zircon and tourmaline became less from E1-2k to N2g, and sedimentary rocks represented by the assemblage of pyrite, barite and limonite also became less. Intermediate-basic igneous rocks represented by the assemblage of leucoxene, rutile and ilmenite and metamorphic rocks represented by epidote became more and more. Electronic microprobe analysis shows that glaucophane and barroisite are existed in Kongdian Formation and the 4th member of Shahejie Formation, and they demonstrate that Western Shandong and Eastern Shandong are all the source regions of the Boxing Sag, and they also indicate that oceanic crust existed before the collision between the Yangtze and North China continent. The fact that Eastern Shandong is the source region of Boxing Sag also indicates that Western Shandong was not high enough to prevent sediment from Eastern Shandong at E1-2k and E2s4. The results of the dating of five detrital zircons of Boxing Sag show Kongdian Formation and the 4th member of Shahejie Formation have the age peaks of 2800Ma and 700-800. It means that Eastern Shandong is the source region of Boxing Sag at early Paleogene and Western Shandong is not high enough to prevent the sediment from Eastern Shandong. The ages of 160-180 and 220-260 Ma, which exist in the Guantao Formation and Paleogene, are common in Eastern Shandong and rare in Western Shandong,and it implied that Western Shandong is a low uplift at 24Ma. The Paleogene strata have almost same age groups, while the Guantao Formation has significant variations of age groups, and this indicates that Boxing Sag and Western Shandong uplift had taken place tremendous changes. The results of apatite fission track in Boxing sag show that three times uplifts happened at the source region at 60 Ma, 45Ma and 15Ma respectively, and the Boxing sag experienced two subsidences at 60Ma, 45Ma and one uplift at 20Ma.