Ecological Engineering 87(2016)91–97
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Ecological
Engineering
j o u r n a l h o m e p a g e :w w w.e l s e v i e r.c o m /l o c a t e /e c o l e n
g
Short
communication
Endophytic culturable bacteria colonizing Lavandula dentata
L.plants:Isolation,characterization and evaluation of their plant growth-promoting activities
S.I.A.Pereira,C.Monteiro,A.L.Vega,P.M.L.Castro ?
CBQF –Centro de Biotecnologia e Química Fina –Laboratório Associado,Escola Superior de Biotecnologia,Universidade Católica Portuguesa/Porto,Rua Arquiteto Lob?o Vital,Apartado 2511,4202-401Porto,Portugal
a r t i c l e
i n f o
Article history:
Received 28April 2015
Received in revised form 5September 2015Accepted 17November 2015
Available online 1December 2015
Keywords:
Endophytic bacteria Diversity
Indol-3-acetic acid Lavender
Plant growth promotion
a b s t r a c t
Aromatic plants such as lavender are stirring the attention of many researchers due to their content in bioactive secondary metabolites that can be used in traditional medicine.However,information regarding naturally occurring lavender associated bacterial endophytes (BE)is limited.To the best of our knowledge,this is the ?rst study which aims to assess the phylogenetic diversity of the culturable endophytic bacteria of Lavandula dentata cultivated under organic management and to evaluate their potential as plant growth promoting (PGP)agents.BE were grouped by random ampli?ed polymorphic DNA and identi?ed by 16S ribosomal RNA gene sequencing.Endophytes were further characterized for the ability to produce several PGP substances,like ammonia,siderophores,indol-3-acetic acid,and hydrogen cyanide and for the ability to solubilize phosphate.Plant cell-wall degrading enzymes were also determined.Densities of BE were higher in roots (log 6.39CFU g ?1fresh weight)than in shoots (log 5.56CFU g ?1fresh weight).Phylogenetic analysis showed that BE were af?liated to two major groups:?-Proteobacteria (50%)and Firmicutes (31.6%)and a small part belonged to ?-(7.9%)and ?-Proteobacteria (10.5%),being Pseudomonas and Bacillus the most highly represented genera.Higher bacterial diversity was found in the lavender roots,with endophytes belonging to 6different genera (Pseudomonas ,Variovorax ,Rhizobium ,Caulobacter,Bacillus and Paenibacillus ),than in shoots where only 3genera (Bacillus ,Pseudomonas and Xanthomonas )were found.Overall,BE showed ability to produce extracellular enzymes and multiple PGP traits,suggesting their potential use as ef?cient bioinoculants in sustainable cultivation of medicinal and aromatic plants.
?2015Elsevier B.V.All rights reserved.
1.Introduction
Lavandula species (Lamiaceae)are endemic to the Mediter-ranean region,Arabian Peninsula,Canary Islands,and India (Upson,2002),being widely cultivated worldwide,due to their economic value for cosmetic and pharmaceutical industries and also for orna-mental purpose (Bakkali et al.,2008;Muyima et al.,2002).Bioactive secondary metabolites such as essential oils are extracted from various aromatic plants and in particular those extracted from Lavandula dentata stand out for their use in popular medicine as antidiabetic,antihypertensive,and antiprotozoal agent (Al-Musayeib et al.,2012),explaining the growing interest of the pharmaceutical industries in these natural compounds.
Lavender species can be propagated by seeds or by woody stem cuttings.The propagation through seeds is usually slow,plants
?Corresponding author.
E-mail address:plcastro@porto.ucp.pt (P.M.L.Castro).
show a large variation in growth and in oil composition,and the genetic uniformity of the resulting plants is not guaranteed (Zuzarte et al.,2010).For these reasons,vegetative propagation is preferred to reproduction by seeds.Propagation by cuttings is fast,does not require special techniques and methods and is less expensive than in vitro micropropagation (Zuzarte et al.,2010).However,the poor rooting capacity of cuttings is often observed.In order to overcome this problem,plant growth regulators like the indole-3-butyric acid (IBA)and indole-3-acetic acid (IAA)are commonly used and showed to be ef?cient in increasing the adventitious root forma-tion and the rooting capacity of cuttings (Bona et al.,2010;Kasem and El-baset,2014).However,chemical application raises environ-mental concerns and increases production costs thus the use of sustainable practices in the production of medicinal and aromatic plants is imperative.
Plant growth promoting rhizobacteria (PGPR)have been used as bioinoculants for improving plant growth in a wide range of environments and crop species (Pereira and Castro,2014a;Pereira et al.,2015).Several authors also reported that PGPR induced root
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92S.I.A.Pereira et al./Ecological Engineering87(2016)91–97
formation in stem cuttings of several aromatic plants,namely in Mentha piperita(Kaymak et al.,2008)and in Lavandula angustifolia (Kasem and El-baset,2014).However,very little is known about the effects of bacterial endophytes(BE)on the rooting capacity of cuttings.
Endophytic bacteria are de?ned as those that colonize the inner tissues of healthy plants without causing symptoms of disease or detrimental effect on their host(Schulz and Boyle,2006).Similarly to rhizobacteria,BE promote plant growth by several mechanisms, including phosphate solubilization,nitrogen?xation,production of siderophores and ammonia and through the production of wide range of phytohormones,such as auxins,cytokinins and gibberellic acids(Pereira and Castro,2014b;Verma et al.,2001).Endophytic bacteria have the advantage of proliferating within the plant tissue thus facing less competition for nutrients and being protected from high-stress environment of the soil(Sturz et al.,2000).Endophytes have been isolated from a wide range of plant species and tissues suggesting their ubiquitous existence in higher plants.However, only a few of all the plants existing on earth have ever been studied relative to their bacterial endophytic pool(Strobel and Daisy,2003), increasing the probability to?nd new and bene?cial endophytes with potential to be applied in biotechnology.The microbiome of medicinal and aromatic plants is extremely important since there are increasing evidences that the spectrum of bioactive metabolites may be related to the activity of associated BE(Emiliani et al.,2014). However,little attention has been paid to the BE from aromatic and medicinal plants and there is a lack of reports in this research area.
The aim of the present study was to isolate and charac-terize BE from tissues(root and shoot)of L.dentata plants growing under organic management.We further intended to assess the phylogenetic diversity and the phenotypic character-ization of the culturable BE;to evaluate their ability to produce plant growth promoting(PGP)substances,such as IAA,ammonia, hydrogen cyanide(HCN),siderophores and extracellular enzymes (lipases,cellulases,proteases,pectinases)and to solubilize phos-phate,in order to indicate BE with potential to be used as bioinoculants in vegetative propagation of aromatic and medicinal plants.
2.Material and methods
2.1.Enumeration and isolation of culturable bacterial endophytes
To isolate representative BE four healthy L.dentata L.plants grown under organic management were collected at random from the“Cantinho das Aromáticas”located in Vila Nova de Gaia, Portugal(GPS–41?07 30.00 N;8?38 40.82 W).Plant surface ster-ilization was performed according to Pereira and Castro(2014b). The success of the surface disinfection process was evaluated by plating100?l of water from the?nal rinse on Trypticase Soy Agar (TSA;Pronadisa)agar medium and incubated at30?C for3days. No bacterial growth was found.
For bacterial enumeration1g of roots and shoots were homog-enized in9ml of sterile phosphate buffered saline(PBS g l?1: Na2HPO4, 1.44;KH2PO4,0.24;KCl,0.20;NaCl,8.00;pH7.4). Serial dilutions were made in duplicate,and0.1ml of each dilution was spread onto TSA and Plate Count Agar(PCA; Merck)media.Plates were incubated for7days at30?C and the number of colony forming units was(CFU)determined. Different bacterial colonies were isolated based on size,morphol-ogy and color,from TSA and PCA media,using a streak-plate procedure.Isolates were puri?ed and further phenotypically characterized.2.2.Random ampli?ed polymorphic DNA(RAPD)and DNA sequencing analysis
Bacterial isolates were grouped according to species sim-ilarity,based on RAPD pro?les produced with primer M13 (5 GAGGGTGGCGGTTCT-3 ).DNA extraction and RAPD analysis were performed according to the methods described in Pereira and Castro(2014b).RAPD ampli?cation products were run on a1.5% agarose gel stained with SYBR Safe(Invitrogen,UK)for135min at80V.RAPD patterns were compared using Bionumerics soft-ware(Applied Maths,St-Martens-Laten,Belgium)and clustered according to their similarities.Isolates displaying unique RAPD pro-?les were subsequently identi?ed by16S rRNA gene sequencing analysis.16S rDNA ampli?cation was performed with universal primers,27F and1492R as previously described by Pereira and Castro(2014b).The PCR products were puri?ed using a GRS PCR& Gel Band Puri?cation Kit(Grisp)and sequencing was performed by Macrogen Inc.(Netherlands).Sequence editing and inspection were performed using BioEdit program7.0version and the sequence similarity search was performed using the EzTaxon sever.For phylogenetic analyses,the sequences were aligned by using the CLUSTALW(Thompson et al.,1994).Tree constructions were per-formed with MEGA5.0,using the neighbor-joining method(Kimura two-parameter distance optimized criteria).The robustness of the phylogenetic tree was con?rmed by using bootstrap analysis based on1000resamplings of the sequences.The16S rRNA sequences of the BE were deposited in GenBank database under accession numbers KP407086to KP407123.Simpson’s(1?D)and Shannon’s (H)diversity indices were calculated based on the percentage of different bacterial genera in both plant compartments.
2.3.Plant growth promotion(PGP)traits
The amount of IAA produced by BE was determined according to Gordon and Weber(1951).Brie?y,an aliquot of500?l of super-natant obtained from bacterial cultures grown in the presence of l-tryptophan(1%)was mixed with350?l of Salper reagent.The absorbance of pink color developed after30min of incubation in dark was read at530nm.The IAA concentration was determined using a calibration curve of pure IAA as a standard.
Detection of siderophore production was carried out by inocu-lating the BE on Chrome Azurol S medium(Schwy and Neilands, 1987).The development of a yellow to orange halo around the colonies after incubation at30?C for72h indicated a positive result for siderophore production.The detection of HCN produc-tion was made by amending nutrient agar with4.4g glycine l?1 and streaking the isolates on this modi?ed agar plates.On the top of each plate,a sterilized?lter paper(Whatman No.1)soaked in 2%sodium carbonate prepared in0.5%picric acid solution was placed.Plates were incubated at30?C for4days after which development of orange to red color indicated HCN production (Ahmad et al.,2008).For assessing the ability to produce ammo-nia,fresh cultures were inoculated into10ml of peptone water and incubated at30?C for48h;following this,0.5ml of Nessler’s reagent(Sigma–Aldrich)was added to each tube and develop-ment of yellow to brown color was considered as a positive result for ammonia production(Cappuccino and Sherman,1992).For phosphate solubilization assay,fresh cultures were inoculated into National Botanical Research Institute Phosphate(NBRIP)medium supplemented with0.5%tricalcium phosphate(Nautiyal,1999). The presence of a clearing halo around bacterial colonies was con-sidered positive for phosphate solubilization.
Bacterial endophytes were screening for their ability to produce extracellular enzymes(proteases,cellulase,pectinase and lipases) according to the methods described in Pereira and Castro(2014b).
S.I.A.Pereira et al./Ecological Engineering87(2016)91–9793
3.Results and discussion
In the last years,several studies have focused their attention on aromatic and medicinal plants due to the presence of distinct bioac-tive secondary metabolites that can be used in traditional medicine. Despite the great interest in medicinal plants,very little is known about the associated BE.To the best of our knowledge,this work is the?rst to assess the phylogenetic diversity of the culturable endo-phytic bacterial populations of L.dentata plants cultivated under organic management and to evaluate their plant growth promoting traits.
In this study,the number of endophytic culturable bacteria var-ied signi?cantly among plant organs.The colony forming units in roots varied between log6.21CFU g?1fresh weight(FW)(for TSA medium)and log6.39CFU g?1FW(for PCA medium)and were higher if compared to shoot where bacterial counts ranged from log5.31CFU g?1FW(for TSA)to log5.56CFU g?1FW(for PCA).Sim-ilar results were obtained for Echinacea purpurea and Echinacea angustifolia where the lowest bacterial counts were determined in the aboveground tissues,while the highest bacterial numbers were detected in roots(Chiellini et al.,2014).A similar trend was observed by Aravind et al.(2008)in black pepper(Piper nigrum) plants.However,several authors reported that population density of endophytes in aromatic and medicinal plants seems to be high-est in aerial than in the belowground tissues.Emiliani et al.(2014) and El-Deeb et al.(2013)showed that leaves had higher numbers of endophytic bacteria than roots in L.angustifolia and Plectranthus tenui?orus plants,respectively.
Based on the distinct colony characteristics a total of56BE were isolated from the plant tissues of L.dentata plants growing under organic management.However,according to RAPD analysis only 38different pro?les were recognized,corresponding to27bacte-rial strains from roots(LR)and11strains from shoots(LS)(Table1). The higher numbers of bacterial counts found in lavender roots, as well as the higher number of endophytes recovered from this tissue,may re?ect the intimate contact of roots with soil rhizo-sphere,which facilitate the entry of bacteria into the root tissues (Kobayashi and Palumbo,2000).In addition,the endophytic bacte-rial population densities reported in the present study were higher than the earlier reports in aromatic and medicinal plants such as P.nigrum(Aravind et al.,2008)and L.angustifolia(Emiliani et al., 2014).These results may be related to the management practices used for the cultivation of lavender plants,since organic manage-ment seems to promote microbial activity in rhizosphere(Reilly et al.,2013),increasing the bacterial pool that can enter into plant tissues.
One isolate of each cluster was chosen for16S rRNA partial gene sequence(Fig.1).As shown in Table1,all BE showed high similarities(98–100%)with their closest related species.Phyloge-netic analyses based on16S rRNA gene sequences showed that BE isolated from lavender tissues belonged to two main groups:?-Proteobacteria(50%)and Firmicutes(31.6%),being a small portion af?liated to?-Proteobacteria(7.9%)and?-Proteobacteria(10.5%). Overall,BE belonged to7different genera,being Pseudomonas (47%)and Bacillus(29%)the better represented in lavender plants (Table1).A similar trend was already described by several authors for other aromatic and medicinal plants.Emiliani et al.(2014) reported that51%of the BE isolated from L.angustifolia tissues belonged to Pseudomonas genus,while Vendan et al.(2010)showed a predominance of the genus Bacillus in ginseng plants.In fact,both genera have been identi?ed as frequently occurring endophytes in several plant species(Hallmann and Berg,2006).
The results also showed that BE colonized differently lavender tissues.In roots,where higher bacterial diversity was found,endo-phytes belonged to6different genera(Pseudomonas,Variovorax, Rhizobium,Caulobacter,Bacillus and Paenibacillus),while in shoots 91%of the endophytic isolates were af?liated to Bacillus and Pseu-domonas and only one isolate belonged to Xanthomonas genus. Some of these genera have already been described in other aro-matic and/or medicinal plants.Cho et al.(2007)isolated63different endophytic strains belonging to13different genera including Bacil-lus and Paenibacillus from the interior of ginseng tissues.Emiliani et al.(2014)also isolated endophytes af?liated to Rhizobium,Pseu-domonas,and Bacillus genera from L.angustifolia plants.However, to the best of our knowledge BE belonging to Caulobacter and Variovorax genera were never detected in lavender plant compart-ments.The differences among the culturable bacterial populations in both lavender organs were also highlighted by the diversity indices calculated based on genera distribution,since Simpson’s and Shannon’s indices were higher in roots(D=0.74and H=1.45, respectively)if compared to shoots(D=0.56and H=0.86,respec-tively).
The distribution of BE in roots and shoots of L.dentata plants may be related to the distinct anatomical and nutritional condi-tions found in both plant compartments,leading to the creation of speci?c ecological niches for endophytic growth.According to Gaiero et al.(2013)endophyte distribution within plants depends on a combination of skills to colonize and the allocation of plant resources.In addition,according to Compant et al.(2010)the secre-tion of cell-wall degrading enzymes by endophytes is an important trait for tissue colonization and bacterial spreading inside plants. In this study,the production of extracellular enzymes such as cellulases,pectinases,proteases and lipases by BE was assessed and the results are shown in Table2.Bacterial endophytes had higher ability to produce pectinases(68%)and proteases(79%) than cellulases(50%)and lipases(58%).In general,endophytic bacteria from shoots showed higher ability to produce plant cell wall-degrading enzymes,especially pectinases and proteases,sug-gesting that endophytic colonization of shoots may be closely related to BE capacity to produce extracellular enzymes.Verma et al.(2001)also reported cellulase and pectinase activities in dif-ferent isolates suggesting their potential for inter-and intracellular colonization.
Rooting of stem cutting is widely used in horticulture for the propagation of aromatic and medicinal plants,however poor roo-ting is often observed(Kasem and El-baset,2014).In order to overcome this problem synthetic auxins are often applied at stim-ulating the adventitious root formation on cuttings(Bona et al., 2010),however due to the continuous increase of environmen-tal footprint it is urgent to?nd eco-friendly alternatives.Although several studies have reported the bene?cial effect of BE and PGPR inoculation in several economically important crops,like maize (Pereira and Castro,2014a),carrot and potatoes(Surette et al., 2003),little information is available for aromatic and medicinal plants.
Screening results of PGP traits of the lavender BE are shown in Table2.All endophytic strains were able to synthesize IAA in the presence of the precursor l-tryptophan,with levels ranging from 7.0to74.7mg l?1.Several BE(21%)produced more than40mg l?1 of IAA and most of them were af?liated to Pseudomonas genus. Bacterial endophytes isolated from ginseng plants also showed ability to synthesize IAA even though at lower levels(Vendan et al., 2010).The application of PGPR in order to minimize the use of syn-thetic auxins was already reported by Kasem and El-baset(2014) and Kaymak et al.(2008)who showed that the inoculation of PGPR in L.dentata cuttings and M.piperina,respectively improved rooting performance.However,so far no studies are available concerning the PGP of BE isolated from aromatic and medicinal plants and their ability to increase stem cutting establishment in soils.Additionally to auxin production,all endophytic strains were able to produce ammonia and siderophores.According to Idris et al.(2004)siderophore production among endophytes may be a
S.I.A.Pereira et al./Ecological Engineering87(2016)91–9795
Fig.1.Neighbor-joining phylogenetic tree based on partial16S rRNA sequences,showing the relationships between sequences of representative strains of bacterial endo-phytes and some of their closest phylogenetic relatives.Bootstrap values are shown at nodes.Acidilobus saccharovorans(NR115208.1)was used as outgroup.Bar indicates 0.05substitutions per nucleotide position.
96S.I.A.Pereira et al./Ecological Engineering87(2016)91–97
Table2
Characterization of bacterial endophytes for multiple plant growth promoting(PGP)traits:NH3,siderophore,IAA,HCN and extracellular enzymes:cellulase,pectinase, protease and lipase production and phosphate solubilization ability.
Strain Closest relatives NH3Siderophores P solubilization IAA(mg l?1)HCN Extracellular enzymes
Cellulase Pectinase Protease Lipase
T20T40T80
LR1-1Pseudomonas brassicacearum
subsp neoaurantiaca
+++20.1±1.84+??????
LR1-2Rhizobium nepotum++?27.8±2.22+??????
LR1-5Pseudomonas moorei+++++46.8±3.39++????+?
LR1-6B Variovorax boronicumulans++?60.5±4.19?+??+++
LR1-7Variovorax soli+++23.1±1.73????+?+
LR1-9Pseudomonas graminis++?51.4±3.58+?+?+??
LR1-11Bacillus cereus biovar toyoi++?25.8±2.99+++++++++
LR1-12Caulobacter vibrioides++?34.1±1.82++++++??
LR1-13Variovorax boronicumulans++?40.2±3.66+???+++
LR1-16Pseudomonas brassicacearum
subsp neoaurantiaca
++?33.4±2.52++???+??
LR2-1Bacillus thuringiensis++? 6.3±0.47++??+++++
LR2-2Pseudomonas k ilonensis++++?26.0±2.26++??+++?+
LR2-3Pseudomonas jessenii++++?17.4±2.55++??+++?+
LR2-4Bacillus cereus biovar toyoi++?25.5±0.70++??++++
LR2-6Bacillus cereus biovar toyoi++++?7.3±1.27+++?++++
LR2-9Pseudomonas punonensis++?9.8±1.10+++++++++
LR2-10Variovorax boronicumulans++?11.7±1.26++?++++++
LR2-11Paenibacillus kribbensis?+?7.3±1.43++++++++?
LR3-1Pseudomonas frederiksbergensis+++8.9±0.55++++++++?+
LR3-2Pseudomonas fuscovaginae+++21.2±1.87+++++++??
LR3-3Bacillus cereus biovar toyoi++++?10.2±0.64+++++++++
LR3-6Pseudomonas fuscovaginae+++?8.3±1.41++++++???
LR3-7Bacillus cereus+++++7.4±1.17+++?+++++?
LR3-8Bacillus aerophilus+++10.2±1.37++++++++
LR3-9Bacillus drentensis+++10.2±1.18++++++???
LR3-10Pseudomonas frederiksbergensis+++35.4±3.95++?++++?+
LR3-11Rhizobium grahamii++?7.0±0.50++?+++++?
LS1-1Pseudomonas graminis+++66.3±8.56+++++?+?
LS1-2Bacillus aryabhattai+++18.4±1.48+++++++???
LS1-4Bacillus cereus biovar toyoi++?19.2±1.36++?++++++
LS1-5Pseudomonas graminis+++14.1±2.42++++++?+?
LS2-1Pseudomonas congelans++?74.7±8.20+++++++??
LS2-2Pseudomonas graminis++?47.9±3.64++++++++?
LS2-3Pseudomonas lutea+++++45.7±3.57++++++?+?
LS3-1Bacillus cereus biovar toyoi++?9.7±1.80+++?++++++
LS3-2Pseudomonas lutea+++17.5±1.94++++++???
LS3-5Pseudomonas koreensis++?8.7±1.13++?++++??
LS3-8Xanthomonas vesicatoria++?18.8±1.50+?+++++++
IAA is expressed as means±SE(n=5–12).(?)negative,(+)positive/weak,(++)intermediate,(+++)strong production.
general phenotype,since endophytes have to compete with plant cells for Fe supply.Several BE also showed ability to solubilize phos-phate,which is of great importance to enhance P availability in soils during the initial colonization.
In this work,several potential human pathogens were isolated from lavender tissues,namely the strains LR1-11,LR2-1,LR2-4,LR2-6,LR3-3,LR3-7,LS1-4and LS3-1that are members of the Bacillus cereus group of bacteria.These BE were found in both plant organs,but with higher incidence in roots.This is an issue of concern,since many genera including Burkholderia,Enterobacter, Bacillus and S tenotrophomonas have been identi?ed as colonizers of the plant rhizosphere but some members can also successfully col-onize human organs and tissues causing diseases(Berg et al.,2005). Consequently,the selection of bacterial endophytes to be applied as bioinoculants in plant cultures should take in consideration the risk of pathogenicity to ensure that they do not inadvertently pose
a threat to human health.
4.Conclusions
Bacterial endophytes seem to colonize differently lavender tissues.Beyond the higher numbers of BE found in the roots,phy-logenetic analysis also showed clear differences between plant organs.In roots,a high diversity of genera was observed and BE were af?liated to Pseudomonas,Variovorax,Rhizobium,Caulobacter, Bacillus and Paenibacillus,while in shoots isolates belonged to Bacil-lus,Pseudomonas and Xanthomonas genera.The BE isolated in this study showed several plant growth promoting traits which suggest their potential for plant growth promotion.The use of such BE as ef?cient bioinoculants may constitute an interesting alternative to the application of chemical compounds,for sustainable cultivation of medicinal and aromatic plants.
Acknowledgments
This work was supported by National Funds through FCT –Fundac??o para a Ciência e a Tecnologia under the project PTDC/AGR/CFL/111583/2009,project PEst-OE/EQB/LA0016/2011 and Fundo Social Europeu(III Quadro Comunitário de Apoio). So?a Pereira and Cristina Monteiro had the support of FCT grants SFRH/BPD/65134/2009and SFRH/BPD/80885/2011,respectively. References
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stocks 存货，库存量 cash sale 现货 purchase 购买，进货 bulk sale 整批销售，趸售 distribution channels 销售渠道 wholesale 批发 retail trade 零售业 hire-purchase 分期付款购买 fluctuate in line with market conditions 随行就市unfair competition 不合理竞争 dumping 商品倾销 dumping profit margin 倾销差价，倾销幅度antidumping 反倾销 customs bond 海关担保 chain debts 三角债 freight forwarder 货运代理 trade consultation 贸易磋商 mediation of dispute 商业纠纷调解 partial shipment 分批装运 restraint of trade 贸易管制 RTA (Regional Trade Arrangements) 区域贸易安排favorable balance of trade 贸易顺差 unfavorable balance of trade 贸易逆差 special preferences 优惠关税 bonded warehouse 保税仓库 transit trade 转口贸易 tariff barrier 关税壁垒
tax rebate 出口退税 TBT (Technical Barriers to Trade) 技术性贸易壁垒 贸易伙伴术语 trade partner 贸易伙伴 manufacturer 制造商，制造厂 middleman 中间商，经纪人 dealer 经销商 wholesaler 批发商 retailer, tradesman 零售商 merchant 商人，批发商，零售商 concessionaire, licensed dealer 受让人，特许权获得者consumer 消费者，用户 client, customer 顾客，客户 buyer 买主，买方 carrier 承运人 consignee 收货人 进出口贸易词汇 commerce, trade, trading 贸易 inland trade, home trade, domestic trade 国内贸易international trade 国际贸易 foreign trade, external trade 对外贸易，外贸import, importation 进口 importer 进口商 export, exportation 出口 exporter 出口商 import licence 进口许口证 export licence 出口许口证
Chapter 1 Passage 1 Human Body In this passage you will learn: 1. Classification of organ systems 2. Structure and function of each organ system 3. Associated medical terms To understand the human body it is necessary to understand how its parts are put together and how they function. The study of the body's structure is called anatomy; the study of the body's function is known as physiology. Other studies of human body include biology, cytology, embryology, histology, endocrinology, hematology, immunology, psychology etc. 了解人体各部分的组成及其功能，对于认识人体是必需的。研究人体结构的科学叫解剖学；研究人体功能的科学叫生理学。其他研究人体的科学包括生物学、细胞学、胚胎学、组织学、内分泌学、血液学、遗传学、免疫学、心理学等等。 Anatomists find it useful to divide the human body into ten systems, that is, the skeletal system, the muscular system, the circulatory system, the respiratory system, the digestive system, the urinary system, the endocrine system, the nervous system, the reproductive system and the skin. The principal parts of each of these systems are described in this article. 解剖学家发现把整个人体分成骨骼、肌肉、循环、呼吸、消化、泌尿、内分泌、神经、生殖系统以及感觉器官的做法是很有帮助的。本文描绘并阐述了各系统的主要部分。 The skeletal system is made of bones, joints between bones, and cartilage. Its function is to provide support and protection for the soft tissues and the organs of the body and to provide points of attachment for the muscles that move the body. There are 206 bones in the human skeleton. They have various shapes - long, short, cube - shaped, flat, and irregular. Many of the long bones have an interior space that is filled with bone marrow, where blood cells are made. 骨骼系统由骨、关节以及软骨组成。它对软组织及人体器官起到支持和保护作用，并牵动骨胳肌，引起各种运动。人体有206根骨头。骨形态不一，有长的、短、立方的、扁的及不规则的。许多长骨里有一个内层间隙，里面充填着骨髓，这即是血细胞的制造场所。 A joint is where bones are joined together. The connection can be so close that no movement is possible, as is the case in the skull. Other kinds of joints permit movement: either back and forth in one plane - as with the hinge joint of the elbow - or movement around a single axis - as with the pivot joint that permits the head to rotate. A wide range of movement is possible when the ball - shaped end of one bone fits into a socket at the end of another bone, as they do in the shoulder and hip joints. 关节把骨与骨连接起来。颅骨不能运动，是由于骨与骨之间的连接太紧密。但其它的关节可允许活动，如一个平面上的前后屈伸运动，如肘关节；或是绕轴心旋转运动，如枢轴点允许头部转动。如果一根骨的球形末端插入另一根骨的臼槽里，大辐度的运动（如肩关节、髋关节）即成为可能。 Cartilage is a more flexible material than bone. It serves as a protective, cushioning layer where bones come together. It also connects the ribs to the breastbone and provides a structural base for the nose and the external ear. An infant's skeleton is made of cartilage that is gradually replaced by bone as the infant grows into an adult. 软骨是一种比一般骨更具韧性的物质。它是骨连结的保护、缓冲层。它把肋骨与胸骨连结起来，也是鼻腔与内耳的结构基础。一个婴儿的骨骼就是由软骨组成，然后不断生长、
贸易方式词汇 stocks 存货，库存量 cash sale 现货 purchase 购买，进货 bulk sale 整批销售，趸售 distribution channels 销售渠道 wholesale 批发 retail trade 零售业 hire-purchase 分期付款购买 fluctuate in line with market conditions 随行就市unfair competition 不合理竞争 dumping 商品倾销 dumping profit margin 倾销差价，倾销幅度antidumping 反倾销 customs bond 海关担保 chain debts 三角债 freight forwarder 货运代理 trade consultation 贸易磋商 mediation of dispute 商业纠纷调解 partial shipment 分批装运 restraint of trade 贸易管制 RTA (Regional Trade Arrangements) 区域贸易安排favorable balance of trade 贸易顺差 unfavorable balance of trade 贸易逆差 special preferences 优惠关税 bonded warehouse 保税仓库 transit trade 转口贸易 tariff barrier 关税壁垒 tax rebate 出口退税 TBT (Technical Barriers to Trade) 技术性贸易壁垒 贸易伙伴术语 trade partner 贸易伙伴 manufacturer 制造商，制造厂 middleman 中间商，经纪人 dealer 经销商 wholesaler 批发商 retailer, tradesman 零售商 merchant 商人，批发商，零售商 concessionaire, licensed dealer 受让人，特许权获得者consumer 消费者，用户 client, customer 顾客，客户 buyer 买主，买方 carrier 承运人 consignee 收货人 进出口贸易词汇 commerce, trade, trading 贸易 inland trade, home trade, domestic trade 国内贸易international trade 国际贸易 foreign trade, external trade 对外贸易，外贸import, importation 进口 importer 进口商 export, exportation 出口
外贸人士必学英语词汇 接下来为大家整理了外贸人士必学英语词汇，希望对你有帮助哦!外贸人士必学英语词汇一：ANER 亚洲北美东行运费协定Asia NorthAmerica EastboundRateB组BAF 燃油附加费Bunker AdjustmentFactorBAF 燃油附加费，大多数航线都有，但标准不一。 B/L 海运提单Bill of LadingB/R 买价Buying RateC组(主要运费已付)CFR 成本加运费(……指定目的港)CFR(cost and freight)成本加运费价C&F(成本加运费)：COST AND FREIGHTC&F 成本加海运费COST AND FREIGHTCIF 成本、保险费加运费付至(……指定目的港)CIF 成本，保险加海运费COST，INSURANCE，FRIGHTCIF(成本运费加保险，俗称“到岸价)：COST INSURANCE AND FREIGHTFOB (离岸价)：FREE ON BOARDCPT 运费付至(……指定目的港)CPT 运费付至目的地Carriage Paid ToCIP 运费、保险费付至(……指定目的地)CIP 运费、保险费付至目的地Carriage and Insurance Paid ToCY/CY 整柜交货(起点/终点)C.Y. 货柜场Container YardCY(码头)：CONTAINER YARDCFS(场)：CARGO FREIGHT STATIONC/D (customs declaration)报关单 C.C.(运费到付)：COLLECTC.C 运费到付CollectCNTR NO. (柜号)：CONTAINER NUMBERC.O (certificate of origin)一般原产地证CTN/CTNS(carton/cartons)纸箱C.S.C 货柜服务费Container Service ChargeC/(CNEE) 收货人ConsigneeC/O 产地证Certificate of OriginCAF 货币汇率附加费Currency Adjustment FactorCFS 散货仓库Container Freight StationCFS/CFS 散装交货(起点/终点)CHB 报关行
汉译英心血管疾病cardiovascular diseases; 脑垂体的功能the function of pituitary; 泌尿道urinary tract; 分子molecule; 动脉artery; 内分泌学endocrinology; 呼吸困难dyspnea; 唾液saliva; 组织学histology; 血液循环blood circulation; 血液学hematology; 生理学physiology; 解剖学anatomy; 女性生殖系统femal reproductive system; 神经细胞nerve cell; 免疫学immunology; 消化不良dyspepsia; 随意肌voluntary muscle; 胚胎学embryology; 心理学psychology; 细胞学cytology; 原生质protoplasm; 细胞膜cell membrane; 细胞核nucleus; 细胞质（浆）cytoplasm; 脱氧核糖核酸deoxyribonucleic acid; 能半渗透的semipermeable; 分子生物学molecular biology; 碳水化合物carbohydrate; 有区别性的differentially; 使…完整intact; 根据according to; 遗传特性hereditary trait; 渗滤diffusion; 转换transaction; 蓝图blueprint; 染色体chromosome; 色素pigment; 排出废液excrete waste fluid; 散开disperse; 脉冲信号impulse; 核糖核酸ribonucleic acid; 损害正常功能impair the normal function; 污染环境pollute environment; 功能失调malfunction; 致病因子causative agents; 易受侵害的人群vulnerable groups; 局部化的感染localized infection; 花柳病venereal disease; 抗原与抗体antigen&antibody; 肌电图electromyogram; 多发性硬化multiple sclerosis; 心电图electrocardiograph; 疾病的后遗症sequelea of disease; 光纤技术fiber optic technology; 造血系统hematopoietic system; 致命的疾病fatal disease; 体液body fluid; 无副作用的治疗hazard-free treatment; 无侵犯的实验检查non-invasive laboratory test; 核磁共振nuclear magnetic resonance; 葡萄糖耐糖实验the glucose-tolerance test; 乐观的预后optimistic prognosis; 超声波检测法ultrasonography; 病史medical history; 随访活动follow-up visit; 营养不良nutritional deficiency; 使细节显著highlight detail; 脑电图electroencephalogram; 缺血的组织blood-starved tissue; 肌纤维muscle fiber; 随意肌voluntary muscle; 消化道alimentary canal; 肌腹fleshy belly of muscle; 横纹肌striated muscle; 肌肉痉挛cramps of muscle; 肌肉收缩muscle contraction; 肌肉附着点attachment of the muscle; 肌肉放松relaxation of muscle; 动脉出血arterial hemorrhage; 止端insertion;起端origion;供血blood supply; 屈肌flexor; 蛋白分子protein molecule; 纤维结缔组织fibrous connective tissue; 伸肌extensor; 意志力willpower; 横切面transverse section; 起搏器pacemaker; 肌萎缩muscle atrophy; 重症肌无力myasthenia gravis; 弥散性局部缺血diffuse ischemia; 常染色体隐性autosomal recessive; 全身性感染systemic infection; 受累的肌肉muscle involved; 显著相关性significant correlation; 神经末梢nerve terminal; 自体免疫反应autoimmune reaction; 神经支配innervation; 肌营养不良muscular dystrophy; 慢性营养不良chronic mulnutrition; 先天性肌病congenital myopathy; 预期寿命life expectancy; 免疫紊乱immunologic derangemant; 发病高峰年龄the peak age of onset; 胸腺肿瘤thymoma; 呼吸肌受累the involvement of respiratory muscle; 感染性肌炎inflammatory myositic; 去神经支配denervation; 矿物质吸收mineral absorption; 机械应力mechanical stress; 骨基质有机部分the organic parts of bone matrix; 青春期早熟premature puberty; 蛋白溶解酶protein-digesting enzyme; 破骨细胞osteoclast; 松质骨spongy bone; 骨折fracture; 不规则骨irregular bone; 骨骼系统skeletal system; 维生素吸收vitamin absorption; 骨钙丧失the loss of calcium from bone; 生长激素growth hormone;
学术英语（医学）课后词组 Unit 1 1.neuron overload 神经过载 2.a typical office visit 典型的诊所就诊 3.DEXA scan DEXA扫描 4.medical practice 行医 5.blood pressure control 血压控制 6.health maintenance 健康保持 7.mammogram report 乳房X线检查报告 8.physical examination 体检 9.side effect of a medication 药物的副作用 10.perpetual panic 永久的恐慌 11.practicing physicians 职业医生 12.transplant field 移植领域 13.medical budget 医疗预算 14.paracetamol tablet 扑热息痛药片 15.childproof cap 防孩子打开的盖子 16.randomized clinical trial 随机临床试验 17.random allocation 随机分配 18.patient prognosis 病人的预后 19.control group 对照组 20.a 10-year follow-up study 10年的跟踪研究 21.a medical ward 内科病房 22.infectious hepatitis 传染性肝炎 23.Severe malaise 身体严重不适 24.bilirubin metabolism 胆红素代谢 25.permanent liver damage 永久的肝损伤 26.exacerbate pathophysiology 加重病理生理状况 27.medical literature 医学文献 28.clinical investigation 临床调查 29.incidence of relapse 复发率 30.clinical epidemiology 临床流行病学 31.strict bed rest 严格的卧床休息 32.hospital stay 住院 33.recurrent jaundice 反复发作的黄疸 34.clinical course 临床病程 35.intravenous morphine 静脉注射吗啡 36.diastolic blood pressure 舒张压37.brain perfusion 大脑血灌输 38.primary care初级保健 39.aorto-coronary arterial bypass主动脉冠状动脉旁路 https://www.doczj.com/doc/119967851.html,rmed treatment decision知情治疗决 41.an international humanitarian group 一个国际人道组织 42.the Red Cross 红十字会 43.the first major relief effort第一次重大援助工作 44.casualty of war 战争中的人员伤亡 45.emergency relief efforts 紧急援助 Unit 2 1.re-emerging/re-emergent/resurgent disease(再现疾病) 2.new flu strain新流感变种 3.antibiotics and vaccine抗生素和疫苗 4.infectious disease传染病 5.emergent/emergent disease新现疾病 6.prevention strategy预防策略 7.bubonic plague腺鼠疫 8.pathogenic microbes病原微生物 9.public heath authority公共卫生机构 10.drug resistance抗药性 11.an course of antibiotic therapy抗生素治疗疗程 12.scarlet fever猩红热 13.the level of virulence毒性水平 14.flu pandemic流感大流行 15.surface antigen 表面抗原 16.genetic shift基因改变 17.neurological complications 神经性并发症 18.waning of immunity免疫力减弱 19.public health infrastructure公共卫生基础设施 20.a malaria case一个疟疾病例 21.swine flu猪流感 22.tuberculosis bacillus结核杆菌 23.the level of morbidity/incident发病
外贸英语词汇全 This model paper was revised by the Standardization Office on December 10, 2020
stocks 存货，库存量 cash sale 现货 purchase 购买，进货 bulk sale 整批销售，趸售 distribution channels 销售渠道 wholesale 批发 retail trade 零售业 hire-purchase 分期付款购买 fluctuate in line with market conditions 随行就市unfair competition 不合理竞争 dumping 商品倾销 dumping profit margin 倾销差价，倾销幅度antidumping 反倾销 customs bond 海关担保 chain debts 三角债 freight forwarder 货运代理 trade consultation 贸易磋商 mediation of dispute 商业纠纷调解 partial shipment 分批装运 restraint of trade 贸易管制 RTA (Regional Trade Arrangements) 区域贸易安排favorable balance of trade 贸易顺差 unfavorable balance of trade 贸易逆差 special preferences 优惠关税 bonded warehouse 保税仓库 transit trade 转口贸易 tariff barrier 关税壁垒 tax rebate 出口退税 TBT (Technical Barriers to Trade) 技术性贸易壁垒 贸易伙伴术语 trade partner 贸易伙伴 manufacturer 制造商，制造厂 middleman 中间商，经纪人 dealer 经销商 wholesaler 批发商 retailer, tradesman 零售商 merchant 商人，批发商，零售商 concessionaire, licensed dealer 受让人，特许权获得者consumer 消费者，用户 client, customer 顾客，客户 buyer 买主，买方 carrier 承运人
医学专业英语 I.Choose the best answer from the four choices marked A, B, C and D (40points) 1.Which of the following suffixes does not mean “pertaining to”? A. -ic B. -al C. -ar D. -our 2.Which of the following combining forms means cell? A. -cyte B. cyt/o C. cel/o D. both a and b 3.Which of the following forms means tissue? A. erythr/o B. immun/o C.hist/o D. vascul/o 4.A- means _____ . A. up B. down C. apart D. without 5.The suffix -itis means _____ . A. infammation B.inflammation C.inflammazed D. instrument 6.The combining form of radi/o means _________ . A. radar B. x-ray C. radio D. both B and C 7.The instrument for viewing is –scope whereas the instrument for cutting is ______ . A. –scopy B. –tome C. –meter D. –graph 8.Which of the following forms refers to “surgical incision o f ?” A. –tomy B. -stomy C. –tome D. –ectomy 9. A record of electric wave occurring in the brain is called _______ . A. electroencephogram B. electromyogram C. electrocardiogram D. electroencephalogram 10.Stethalgia means the pain of chest, which can also be expressed in ____ . A. stethodynia B. thoracalgia C. thoracodynia D. all of the above 11.Which of the following forms means fungus? A. my/o B. myel/o C. myc/o D. none of the above 12.The combining form meaning “the skin” is _______ . A. dem/o B. derm/o C. dermat/o D. both B and C 13.Which of the following combining forms means blood vessels ? A. vascul/o B. angi/o C. chromat/o D. Both A and B 14.The term meaning “pertaining to secretion” is _______ . A. crinogenic B. endocrine C. crinologic D. endocardial 15.The disease of having too many white blood cells is referred to as ________ . A. leukocyte B. leukemia C. leukogenesis D. erythrocyte 16.Inflammation of liver is known in English as ________ . A. bronchitis B. pneumonitis C. gastritis D. hepatitis 17.-plasm means ______ . A. treatment B. knowledge C. diagnosis D. growth 18.Mammo/o refers to ______ . A. mother B. thymus gland C. thyroid gland D. breast 19.Which of the following does not mean “within or in”? A. en- B. endo- C. intra- D. none of the above 20._____ means surgical repair of an organ. A. phag/o B. –plasm C. –plasty D. –pathy
学术英语医学课后问题 答案 HEN system office room 【HEN16H-HENS2AHENS8Q8-HENH1688】
U n i t 1 1、Some factors that may lead to the complaint: ·Neuron overload ·Patients* high expectations ·Mistrust and misunderstanding between the patient and the doctor 2、Mrs. Osorio’s condition: ·A 56-year-old woman ·Somewhat overweight ·Reasonably well-controlled diabetes and hypertension ·Cholesterol on the high side without any medications for it ·Not enough exercises she should take ·Her bones a little thin on her last DEXA scan 3、 Good things: ·Blood tests done ·Glucose a little better ·Her blood pressure a little better but not so great Bad things:·Cholesterol not so great ·Her weight a little up ·Her bones a little thin on her last DEXA scan 4 4、The situation: ·The author was in a moderate state of panic: juggling so many thoughts about Mrs. Osorio’s conditions and trying to resolve them all before the clock ran down. ·Mrs. Osorio made a trivial request, not so important as compared to her conditions. ·Mrs. Osorio seemed to care only about her “innocent — and completely justified —request”： the form signed by her doctor. ·The doctor tried to or at least pretended to pay attention to the patient while completing documentation. 5、Similarities:
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NECK SEAM颈圈 NET WT净.重 NON-FUSIBLE INTERLING非粘朴 NON-WOVEN FABRIC非织布 / 无纺布 NOTCH扼位 OFF PRESSING终烫 OIL STAIN油污 ONE PIECE DOUBLE FOLDED BELT-LOOP一片双折裤耳ONE-PIECE DRESSxx OPEN SEAM开骨 OPERATION BREAK DOWN分工序 OUT-SEAM外骨 OUT-SEAM PKT侧.骨袋 OVERALL工作服 OVERALLS吊带裤 OVERLAP重叠 OVERLAPPING A FEW STITCHING驳线OVERLOCK & BLIND-STITCH折挑 OVERLOCK W/ 5 THREADS五线锁边OVERLOCK WITH 5 THREADS五线锁边OVERTIME WORKING加班工作
Unit1 1、Some factors that may lead to the complaint: ·Neuron overload ·Patients* high expectations ·Mistrust and misunderstanding between the patient and the doctor 2、Mrs. Osorio’s condition: ·A 56-year-old woman ·Somewhat overweight ·Reasonably well-controlled diabetes and hypertension ·Cholesterol on the high side without any medications for it ·Not enough exercises she should take ·Her bones a little thin on her last DEXA scan 3、Good things: ·Blood tests done ·Glucose a little better ·Her blood pressure a little better but not so great Bad things: ·Cholesterol not so great ·Her weight a little up ·Her bones a little thin on her last DEXA scan 4 4、The situation: ·The author was in a moderate state of panic: juggling so many thoughts about Mrs. Osorio’s conditions and trying to resolve them all before the clock ran down.
【外贸必备】外贸术语中英对照 在国际贸易过程中，我们会遇到很多相关行业术语，英文不太好的小伙伴们总是傻傻搞不清，今天我帮大家整理了一些外贸术语中英文介绍，英文不好也不用怕了。美国海关申报费 AMS fee 海运燃油附加费 BAF Bunker Adjustment Fee 并单费combinaton bill of lading fee 出口报关费export customs declaration fee 订舱费booking fee 包干费all in charges 保险费insurance premium 保证金deposit 仓储超期费warehouse overdue fee 仓储费warehouse storage fee 产地证certificate of original 卡车费frucking fee 冲港费special wharf charge
出口操作费export handing charges 重签费reseal fee 拆箱费devanning fee 查验费customs inspection fee 操作费handing charges 超重费over weight charges 堆存费overtime strage charges 目的港手续费destination delivery charges 代垫费用other charges 电放费surrender charges 到付佣金collect commission 地面服务费ground agent service fee 文件费document fee 待时费waiting fee 倒箱费exchange container fee 附加费additional charges
cardiovascular diseases; 脑垂体的功能the function of pituitary; 泌尿道urinary tract; 分子molecule; 动脉artery; 内分泌学endocrinology; 呼吸困难dyspnea; 唾液saliva; 组织学histology; 血液循环blood circulation; 血液学hematology; 生理学physiology; 解剖学anatomy; 女性生殖系统femal reproductive system; 神经细胞nerve cell; 免疫学immunology; 消化不良dyspepsia; 随意肌voluntary muscle; 胚胎学embryology; 心理学psychology; 细胞学cytology; 原生质protoplasm; 细胞膜cell membrane; 细胞核nucleus; 细胞质(浆)cytoplasm; 脱氧核糖核酸deoxyribonucleic acid; 能半渗透的semipermeable; 分子生物学molecular biology; 碳水化合物carbohydrate; 有区别性的differentially; 使…完整intact; 根据according to; 遗传特性hereditary trait; 渗滤diffusion; 转换transaction; 蓝图blueprint; 染色体chromosome; 色素pigment; 排出废液excrete waste fluid; 散开disperse; 脉冲信号impulse; 核糖核酸ribonucleic acid; 损害正常功能impair the normal function; 污染环境pollute environment; 功能失调malfunction; 致病因子causative agents; 易受侵害的人群vulnerable groups; 局部化的感染localized infection; 花柳病venereal disease; 抗原与抗体antigen&antibody; 肌电图electromyogram; 多发性硬化multiple sclerosis; 心电图electrocardiograph; 疾病的后遗症sequelea of disease; 光纤技术fiber optic technology; 造血系统hematopoietic system; 致命的疾病fatal disease; 体液body fluid; 无副作用的治疗hazard-free treatment; 无侵犯的实验检查non-invasive laboratory test; 核磁共振nuclear magnetic resonance; 葡萄糖耐糖实验the glucose-tolerance test; 乐观的预后optimistic prognosis; 超声波检测法ultrasonography; 病史medical history; 随访活动follow-up visit; 营养不良nutritional deficiency; 使细节显著highlight detail; 脑电图electroencephalogram; 缺血的组织blood-starved tissue; 肌纤维muscle fiber; 随意肌voluntary muscle; 消化道alimentary canal; 肌腹fleshy belly of muscle; 横纹肌striated muscle; 肌肉痉挛cramps of muscle; 肌肉收缩muscle contraction; 肌肉附着点attachment of the muscle; 肌肉放松relaxation of muscle; 动脉出血arterial hemorrhage; 止端insertion;起端origion;供血blood supply; 屈肌flexor; 蛋白分子protein molecule; 纤维结缔组织fibrous connective tissue; 伸肌extensor; 意志力willpower; 横切面transverse section; 起搏器pacemaker; 肌萎缩muscle atrophy; 重症肌无力myasthenia gravis; 弥散性局部缺血diffuse ischemia; 常染色体隐性autosomal recessive; 全身性感染systemic infection; 受累的肌肉muscle involved; 显著相关性significant correlation; 神经末梢nerve terminal; 自体免疫反应autoimmune reaction; 神经支配innervation; 肌营养不良muscular dystrophy; 慢性营养不良chronic mulnutrition; 先天性肌病congenital myopathy; 预期寿命life expectancy; 免疫紊乱immunologic derangemant; 发病高峰年龄the peak age of onset; 胸腺肿瘤thymoma; 呼吸肌受累the involvement of respiratory muscle; 感染性肌炎inflammatory myositic; 去神经支配denervation; 矿物质吸收mineral absorption; 机械应力mechanical stress; 骨基质有机部分the organic parts of bone matrix; 青春期早熟premature puberty; 蛋白溶解酶protein-digesting enzyme; 破骨细胞osteoclast; 松质骨spongy bone; 骨折fracture; 不规则骨irregular bone;