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Mongolian Medicine echinops prevented postmenopausal osteoporosis and induced ER/AKT/ERK pathway in BMSCs Yan Liu 1,2, § , Xiongyao Wang 3, § , Hong Chang 2 , Xiaoming Gao 2 , Chongyang Dong 3 , Zimu Li 3 Jingtao Hao 3 , Jiuhe Wang 4 , Qiaoling Fan 1, * 1 School of Basic Medical Science, Nanjing University of Chinese Medicine, Nanjing, China; 2 Department of Traditional Chinese Medicine, Affiliated Hospital of Inner Mongolia Medical University, Hohhot, China; 3 College of Traditional Chinese Medicine, Inner Mongolia Medical University, Hohhot, China; 4 Department of Cardiology, Inner Mongolia Autonomous Rengion Hospital of Traditional Chinese Medicine, Hohhot, China. 1. Introduction Osteoporosis (OP) is a metabolic bone disease characterized by low bone mass and the destruction of the microstructure of bone tissue, leading to increased bone fragility and easy to fracture ( 1,2 ). OP is the sixth most common chronic diseases in humans ( 3 ). It was divided into two types: primary OP and secondary OP ( 4,5 ). Primary OP can be divided into two subtypes, namely Type I and Type II. Type I is postmenopausal osteoporosis (PMOP), which occurs in postmenopausal women ( 6,7 ). Type II is senile osteoporosis, most 200 million people worldwide suffer from osteoporosis, of which postmenopausal women account for 1/3 ( 8,9 ). The overall incidence of OP in Chinese population over 60 is 22.6%, with 15% for males and 28.6% for females, with a trend of increasing year by year ( 10 ). The incidence of OP in the United States is also quite high, about 20,000 cases of OP fractures, and 65,000 cases Summary effective in decreasing the risk of osteoporosis. Mongolian medicine echinops prevents osteoporosis, but its mechanism remains unclear. In this study, we explored the mechanism underlying echinops prevents and treats postmenopausal osteoporosis. Osteoporosis model was established by ovariectomy in rats. Rats were treated to Echinops (16.26, 32.5, or 65 mg/ kg/day) by oral gavage for 3 months. Bone mineral density (BMD) was detected by micro- CT detection of left proximal medial metaphyseal tibia. Hematoxylin and eosin (H&E) and toluidine blue O staining were also performed. Serum levels of E2, ALP and testosterone treated with echinops-containing serum. Estrogen receptors (ER) including ERα and ERβ in bone specimens and BMSCs were detected by qRT-PCR. Cell viability and colon formation of BMSCs were detected. Expressions of ERα, ERβ, AKT, p-AKT, ERK, and p-ERK in BMSCs were detected by western blot. Results showed that echinops significantly increased trabecular interconnectivity, thickness of trabeculae, and connection of trabecula. Echinops significantly increased BMD and E2, but significantly reduced ALP and testosterone in dose- Echinops enhanced cell viability and ability of colony formation of BMSCs, and increased ERα, ERβ, p-AKT, and p-ERK. Thus, Mongolian echinops reduced bone loss and delayed the occurrence and development of osteoporosis, and increased ERα, ERβ, p-AKT, and P-ERK in BMSCs. These results provide experimental basis for clinical prevention and treatment of postmenopausal osteoporosis by echniops. Keywords: Osteoporosis, echinops, ERα, ERβ, AKT/ERK pathway DOI: 10.5582/bst.2018.01046 www.biosciencetrends.com BioScience Trends Advance Publication 2018. § These authors contributed equally to this work. * Address correspondence to: Dr. Qiaoling Fan, School of Basic Medical Science, Nanjing University of Chinese Medicine, 138 Xianlin Road, Qixia District, Nanjing, Jiangsu 210023, China. E-mail: njfanql@163.com; QLFan1803@yeah.net Original Article Advance Publication died due to OP each year ( 11 ). PMOP occurs in 5 to 10 years after menopause in women, most of whom have an increased bone turnover rate, due to fluctuations or gradually reduced in the level of estrogen. In post-menopausal 5-7 years, women lose about 20% of the bone mass ( 12 ). Although most osteoporosis does not directly cause death, its greatest risk is fractures, with high morbidity and disability ( 6,11 ). Estrogen deficiency caused by postmenopausal ovarian hypofunction is recognized as an important cause of postmenopausal osteoporosis ( 13-15 ). Estrogen replacement therapy is the preferred method of treatment for PMOP, which can enhance bone mineral density (BMD) and systemic bone mineral content, and effectively treat postmenopausal osteoporosis ( 16-18 ). However, the long-term use of estrogen increases the risk of breast cancer, endometrial cancer, cardiovascular accident and vascular embolism ( 19 ). In recent years, the prevention and treatment of PMOP by traditional Chinese medicine is attracting more and more attention ( 20,21 ). It is of great significance to find an estrogen replacement medicine for the prevention and treatment of postmenopausal osteoporosis in traditional medicine. Mongolian medicine echinops was introduced in the canon of Mongolian Medical "Wisdom Ancientmirror". It functions as strengthening bone, reuniting bone, and callus ( 22 ). In recent year, it was showed that echinops decreased the serum level of bone Glp protein and inhibited osteoporosis in ovariectomized (OVX) rats ( 23 ). Post-surgery 90 days, the OVX rats were filled the stomach with echinops for 90 days and then the serum level of alkaline phosphatase (ALP) were significantly increased and serum level of interleukin-1 was significantly reduced ( 24 ). After successes of the osteoporosis, OVX rats were feed 90 days and then filled the stomach with echinops for 90 days, then BMD and the maximum deflection of bone were increased compared with OVX rats ( 25 ). These results implied that Mongolian echinops can inhibit the bone absorption and promote the bone formation, decreasing bone turnover, reducing bone loss, delaying the occurrence and development of PMOP. However, the mechanism underlying echinops prevents and treats PMOP is still unclear. In this study, we aimed to explore the roles of estrogen receptors (ER; ERα and ERβ), p-AKT, and p-ERK in BMSCs during echinops prevents and treats postmenopausal osteoporosis. 2. Materials and Methods 2.1. Animals and treatments A total of 84 SPF healthy female Wistar rats (250 ± 20 g, 4 months) were purchased from the animal research center of Inner Mongolia University, China (certification number: SCXK (Mongolia) 2012-0001). The rats received ad libitum access to standard chow pellets and water in 24°C, 50-60% humidity. This study was approved by animal ethics committee of Inner Mongolia University. After 7 days in new environment, the rats were anesthetized by intraperitoneal injection of 40 mg/ kg pentobarbital sodium (P3761, SIGMA-ALDRICH, USA), shaved off the hair on the bilateral dorsal regions for OVX surgery ( 26 ). The ovaries were exposed by a 2 mm incision and resected with surgical scissors. Then, other exposed tissues were repositioned and incision was sutured with 3.0 silk threads in a routine fashion. Intraperitoneal injection of penicillin was administrated. Rats in sham groups was incision without ligation of ovaries artery. Post-surgery 3 months ( 26 ), animals were treated to echinops (16.26, 32.5, or 65 mg/kg/day) by oral gavage for 3 months as previously reported ( 25 ). Echinops was prepared as below: weigh 500 g of dried echinops at 75°C, mixed with 10-time water, decocted for 1 h and filtrated; the slag was decocted with same volume water and filtrated again; the two filtration solutions were collected and concentrated to 500 ml to obtain 1g/mL echinops stock solution. The rats in OVX and sham groups received PBS daily. E2 treatment (E2758, SIGMA-ALDRICH) was set as positive control. After 3 months, the rats were euthanized by intraperitoneal injection of 40 mg/kg pentobarbital sodium, whole blood was collected from the heart through cardiac puncture, and the femur medial malleolus specimens were selected at 1 mm under the epiphyseal plate. After centrifuged at 3,000 rpm for 10 min, serum samples were collected, filtered with 0.22 μm filter, and stored at -20°C for subsequent experiments. 2.2. Hematoxylin and eosin (H&E) and toluidine blue O staining The femur bones were fixed in 10% neutral buffered formalin solution for 48 h, dehydrated in graded ethanol (70-100%, cleared in xylene, embedded in paraffin, and sectioned into 5 μm. For H&E staining, sections were stained with hematoxylin for 3-8 min and eosin for 1-3 min. For toluidine blue O staining, sections were rinsed in toluidine blue O solution for 1 min. The images were observed by Olympus BX51 light microscopy (Olympus, Japan). 2.3. Micro-computed tomography (micro-CT) detection Micro-CT of left proximal medial metaphyseal tibia were acquired using Scanco Μct35 scanner (Scanco, Switzerland) at 70 KVp, 114 μA for 800 ms. Bone mineral density was evaluated based on the micro-CT results. 2.4. Serum levels of E2, ALP, and testosterone After centrifuged at 3,000 rpm for 10 min, serum samples P2 P3 were seeded in 6-well plates at 8 × 10 4 cells/well with echinops-containing serum for 13 days. Then, cells were fixed with 4% paraformaldehyde and stained with 0.1% crystal violet. 2.8. Western blotting Protein was extracted using RIPA (Beyotime, China) with PMSF (1:100) at 4℃ for 30 min, and quantified by BCA assay (#23227, Thermo, USA). 30 µg protein was separated by 8%SDS-PAGE and transferred to PVDF membranes (IPVH00010, Millipore, USA). The membrane was incubated with 5% non-fat milk for 30 min, and incubated with primary antibodies ERα (1:2,000), ERβ (1:4,000), AKT (1:1,500), p-AKT (1:1,000), ERK (1:1,000), p-ERK (1:2,000), and GAPDH (1:10,000) antibody at room temperature for 1 h, and then incubated with HRP goat anti-rabbit IgG secondary antibody (1:20,000, BOSTER, China) at room temperature for 40 min. The blots were detected using Immobilon Western CHEMILUM HRP Substrate (WBKLS0500, Millipore, USA), and light-producing reactions are captured with X-ray film. 2.9. Statistical analysis Data were expressed as mean ± standard deviation, and compared using one-way analysis of variance (ANOVA) followed by Tukey's multiple comparison test. Statistical analyses were performed using the SPSS 10.0 software (SPSS, USA). Significance was considered at p 3. Results 3.1. Echinops increased BMD and E2, but reduced ALP and testosterone in dose-dependent manners BMD at 90 days post-treatment was analyzed by micro-CT (Figure 1A). Compared with sham group, a significant reduction in BMD of cortical bone in OVX group was observed ( p ) not significantly increased the trabecular BMD, whereas 32.5 mg/kg ( p )( p ) significantly increased the BMD of trabecular bone in comparison to the untreated OVX rats. The rats in OVX group showed lower level of E2 ( p )ALP ( p 0.01, Figure 1C) and testosterone ( p 1D) than sham group. 16.26 mg/kg echinops not significantly changed the levels of E2 and testosterone, but significantly inhibited ALP level ( p ) comparison to the OVX group. 32.5 mg/kg and 65 mg/kg echinops significantly increased E2 level but decreased levels of ALP and testosterone in OVX rats (Figure 1B- D). There was not significant difference in ALP level between 32.5 mg/kg and 65 mg/kg treatment groups (Figure 1C). were collected and stored at -20°C until enzyme-linked immunosorbent assay (ELISA) detection. The levels of E2 (CSB-E05108h, CUSABIO), ALP (A059-2, Nanjing Jiancheng Bioengineering Institute), and testosterone (05099h, CUSABIO) were determined by commercially ELISA kits according to manufacturer's instruction using an Multiskan microplate reader (Thermo, USA). 2.5. qRT-PCR detection Total RNA from bone specimens were extracted using Trizol (Takara, Japan). The RNA quality and quantity were examined using Nanodrop 1000 spectrophotometer (NanoDrop, USA). RT reaction was performed using Bestar qPCR RT kit (ABI, USA) according to manufacturer's instruction on ABI9700 PCR system (ABI, USA). The PCR reaction was performed using DBI Bestar ® SybrGreen qPCRmasterMix (ABI, USA) on Stratagene Mx3000P Real time PCR system (Agilent, USA) according to manufacturer's instruction. The primer was listed below (5'-3'): R-GADPH Forward CCTCGTCTCATAGACAAGATGGT, reversed GGGTAGAGTCATACTGGAACATG; ERα Forward AAGAAGAATAGCCCCGCCC, reversed GCCAGGTTGGTCAATAAGCC; ERβ Forward ATGCCCTGGTCTGGGTGAT, reversed CCCCGAGATTGAGGACTTGT. Glyceraldehyde 3-phosphate dehydrogenase (GAPDH) was used as internal control. Relative expression was calculated using 2 method and normalized to sham group. 2.6. Bone marrow-derived bone marrow stem cell (BMSC) isolation and flow cytometry identification The rat femur was rinsed in PBS containing 1% penicillin-streptomycin. After removal of both ends in joints, the bone was rinsed in DMEM with low glucose by a syringe until the bone pale. The mediums were collected and centrifuged at 800 rpm for 5 min. Cells were resuspended into DMEM with low glucose and cultured in incubator at 37°C, 5%CO 2 . After 48 h, cells were stained with primary antibodies of CD29 (ab179471, Abcam, USA), CD90 (ab216449, Abcam), CD45 (ab10558, Abcam), CD11b (ab128797, Abcam), and FITC-conjugated secondary antibody (ab6717, Abcam) in the darker for 30 min. The CD29, CD90, CD45, CD11b positive cells were analyzed using Epics-XL II flow cytometry (Beckman Coulter, USA). 2.7. Cell viability and colon formation Cell viability was detected with a cell counting-8 kit (CCK-8, Beyotime, China). Cells (5x105 cells) were seeded in 96-well plates and incubated with echinops- containing serum for 24, 48 and 72h. Then, CCK-8 was added and the absorbance was detected at 450 nm on microplate reader. For colon formation assay, cells P4 Thus, echinops increased BMD and level of E2, but decreased levels of ALP and testosterone in concentration- dependent manners. 3.2. Echinops induced expression of ERα and ERβ To detect the role of echinops on ERα and ERβ expression, qRT-PCR assays were performed (Figure 2). Compared with sham group, a significant decrease in ERα and ERβ mRNA expressions were shown in OVX rats (Figure 2A and B). Echinops administrated at 16.26, 32.5 and 65 mg/kg significantly increased ERα and ERβ expressions in comparison to OVX group, showing a dose-dependent manner (Figure 2A and B). 3.3. Echinops inhibited osteopenia induced by OVX After surgery for 90 days ( 26 ), rats in sham and OVX group were treated with echinops (32.5 mg/kg) or E2 for 90 days. Then, rats were euthanized and bones were collected. HE and toluidine blue O stainings showed a typical osteopenia with widened intertrabecular spaces, loss of trabecular bone thickness and interconnectivity in OVX group, compared with sham group (Figure 3). Echinops or E2 treatment significantly increased trabecular interconnectivity, thickness of trabeculae, and connection of trabecula, compared with OVX group, suggesting echinops inhibited the osteopenia induced by OVX (Figure 3). 3.4. Isolation and identification of BMSCs To explore the mechanism in echinops treated osteoporosis, BMSCs were isolated and identified by flow cytometry of CD29, CD90, CD45, and CD11b. There were 90.6% isolated-cells positively expressed CD29 (Figure 4A), 93.8% positively expressed CD90 (Figure 4B), but only 4.5% positively expressed CD45 (Figure 4C), only 1.3% positively expressed CD11b (Figure 4D). These results suggested the isolated cells were almost BMSCs. 3.5. Echinops enhanced cell viability and ability of colony formation of BMSCs To evaluate role of echinops in cell proliferation of BMSCs, cell viability (Figure 5A) and colony formation (Figure 5B and C) were performed. Echinops-containing serum (65 mg/kg) significantly increased cell viability (Figure 5A). Moreover, echinops-containing serum significantly increased colony formation of BMSCs in a dose-dependent manner ( p mg/kg, and p )These Figure 3. Echinops inhibited osteopenia induced by OVX. After surgery for 90 days, rats were treated with echinops (32.5 mg/ kg) or E2 for 90 days. HE and toluidine blue O staining were performed. 200X; arrow heads: loss of interconnectivity; arrow: trabecular bones. Figure 1. Micro-CT detection of bone mineral density (BMD) and Elisa detection of E2, ALP, testosterone levels. After surgery for 90 days and treatment for 90 days, (A) micro- CT was performed. Serum E2 (B) , ALP (C) , and testosterone (D) levels were detected by ELISA. Low: 16.26 mg/kg; middle: 32.5 mg/kg; high: 65 mg/kg. * p ** p *** p n.s: not significant by ANOVA test. Figure 2. Effects of echinops on ERα and ERβ expression. After surgery for 90 days and treatment for 90 days, qRT- PCR detection of ERα mRNA (A) and ERβ mRNA (B) . Low: 16.26 mg/kg; middle: 32.5 mg/kg; high: 65 mg/kg. * p ** p *** p ANOVA test. P5 results suggested echinops-containing serum induced cell proliferation of BMSCs in a dose-dependent manner. 3.6. Echinops increased ERα, ERβ, p-AKT, and P-ERK in BMSCs To explore the mechanism in echinops treated osteoporosis, expressions of ERα, ERβ, AKT, p-AKT, ERK, and p-ERK in BMSCs after echinops-containing serum treatment were examined (Figure 6). After treatment of Echniops-containing serum, ERα and ERβ levels were significantly increased compared with control BMSCs (Figure 6A and B). Moreover, the phosphorylations of AKT and ERK were significantly induced in a dose-dependent manner (Figure 6C and D). 4. Discussion In this study, it was demonstrated that echinops functions like estrogen. It can effectively prevent and treat PMOP. Administrating echinops to ovariectomy-induced PMOP model, the BMD and serum level of E2 were increased, serum levels of ALP and testosterone were decreased. Echinops induced expression of ERα and ERβ in OVX rats. The mechanism in echinops prevented PMOP was explored by treating isolated BMSCs with echinops-containing serum. Echinops-containing serum significantly increased cell viability and colony formation of BMSCs, and increased ERα, ERβ, p-AKT, Rats are the most used model animal in the studies of osteoporosis so far ( 14,27 ). After ovariectomy in female rats, bone turnover accelerated, bone loss and bone strength decreased, which was similar to that of people after menopause ( 28 ). Using OVX model, it is Figure 4. Identification of BMSCs isolated from rats. Flow cytometry detection of (A) CD29, (B) CD90, (C) CD45 and (D) CD11b on isolated cells were performed. Figure 5. Effects of echinops-containing serum on cell viability and colony formation. BMSCs were treated with Echniops-containing serum that collected from rats administrated with 16.26 (low), 32.5 (middle), and 65 (high) mg/kg echinops. (A) Cell viability. (B, C) Colony formation. * p ** p *** p vs . control. Figure 6. Effects of echinops-containing serum on expressions of ERα, ERβ, AKT, p-AKT, ERK, and P-ERK in BMSCs. BMSCs were treated with Echniops-containing serum that collected from rats administrated with 16.26 (low), 32.5 (middle), and 65 (high) mg/kg echinops. (A) ERα mRNA. (B) ERβ mRNA. (C) ERα, ERβ, AKT, p-AKT, ERK, and p-ERK. (D) Quantification of western bolts. * p 0.05, ** p 0.01 vs . control. P6 easy to observe the effect of aging on bone tissue, the distribution and reconstruction of cancellous bone in rats and trabecular bone reconstruction of lamellar bone that similar to human ( 28 ). In the present study, a PMOP animal model was established in ovariectomized rats. The ovaries were artificially removed and the estrogen deficiency was induced in rats. The ERα and ERβ levels were also decreased. After 3 months, osteoporosis model was successfully replicated in OVX rats as previously described ( 26 ). Estrogen is recognized drug in the prevention and treatment of PMOP, and diethylstilbestrol (E2) is a synthetic non-steroidal estrogen which can produce pharmacological and therapeutic effect similar to natural estradiol, and significantly reduce the ovariectomy- induced high bone turnover and -reduced bone resorption ( 29,30 ). Therefore, this study selected E2 as a positive control drug to validate the mechanism of Mongolian medicine echinops in PMOP treatment. Similar effects to E2 was demonstrated in this study. After menopause, women's estrogen levels decreased significantly, estrogen through the estrogen receptor (ER) directly effect on the osteoblast and osteoclasts, lead to imbalance of bone resorption and bone formation, resulting in reduced bone mass and BMD, increased bone fragility and the occurrence of osteoporosis ( 31,32 ). The decrease of estrogen level and the decrease of the expression of ER in bone tissue are one of the most important pathogenesis. Estrogen can directly stimulate osteoblasts formation, inhibit osteoclasts activity, and regulate and control the balance of bone formation and bone resorption through ER. ER is expressed in both osteoblasts and osteoclasts ( 33,34 ). Osteoclasts is a very active metabolic giant multinucleated cells, recruited in the bone surface, and played important role in bone resorption and formation of lacuna through release of enzyme and acidic substances such as ALP to dissolve the bone matrix ( 34,35 ). Osteoblasts can synthesize the basic bone material, induces the formation of bone ( 33 ). The combination of estrogen and ER in osteoclasts induces the apoptosis of osteoclasts and osteoclasts precursors, reduces the number of osteoclasts. On the other hand, the combination of estrogen and ER in osteoclasts inhibited the recruitment and differentiation of osteoclasts precursors. In addition, estrogen regulated by osteoprotegerin/osteoprotegerin ligand (OPG/OPGL) system ( 36 ). The decline in estrogen levels results in a dysregulated ratio of osteoprotegerin/osteoprotegerin ligands leading to PMOP ( 37 ). It was demonstrated that echinops decreased the serum level of bone Glp protein, interleukin-1, but increased serum level of ALP, increasing BMD, and inhibiting osteoporosis in ovariectomized (OVX) rats ( 23-25 ). Activation of ERK/PI3K plays important role in ER-mediated cell proliferation ( 38 ). Daidzein stimulated osteogenesis was mediated by both ERα and ERβ, and activation of ERK/PI3K pathway ( 39 ). Consistently, the findings in this study showed echinops reduced ALP and testosterone serum levels in OVX rats, increased BMD and inhibited osteoporosis. In isolated BMSCs, echinops induced cell proliferation, increased ERα, ERβ, p-AKT, and P-ERK. This might associate with the enhancement of osteoblasts differentiation from BMSCs, inhibiting bone absorption and promote bone formation. In conclusion, Mongolian echinops reduced bone loss and delayed the occurrence and development of PMOP, and increased ERα, ERβ, p-AKT, and p-ERK in BMSCs. 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Advanced ImmunoChemical

Advansta

AFFILAND

Affiniti

Affinity Biosciences

AffinityImmuno

Agilent Technologies

Agrisera

Aldevron

Aldrich

Allelebiotech

AllStar Scientific

Alomone

Alomone Labs

ALPCO

Alpha Diagnostics

American Research Products

Ameritech Biomedicines

Amgen

Ampersand Biosciences

Amsbio

AmyJet Scientific

AnaSpec

Anawa

Anbo Biotechnology

Ancell

Angiobio

Aniara

Anogen

Antibodies Inc

Antibodies-online

Antigenix America

APES

Applichem

Argene

Arigo

Astra Biotech GmbH

ATCC

ATGen

Athenaes

Athens Research and Technology

Atlas Antibodies

Atsbio

Aurion

Austral Biologicals and Biogenesis Ltd

Autogenbioclear

Aves Labs

Aviscera Bioscience

Aviva Systems Biology

Bachem

Badrilla

Base Pair Biotechnologies

BD Biosciences

Beckman Coulter

BEI Resources

Bergisch

Bertin Pharma

Bethyl

Bindingsite

Bio X Cell

Bio-Rad

Bio-Rad

Bio-Rad

Bioacademia

Biocare Medical

BioCarta

Biocat

Biochain

Biocheckinc

Biochem

Biocytex

Biogen

Biogenex

Bioke

Biolegend

BioLegend

BioLogo

Biomacromolecules

Biomaterial

Biomatik

Biomedica Medizinprodukte GmbH & Co KG

Bioporto

Biorbyt

Bioscience

Biosensis

Bioss

Bioss

Biotech

Biotium

Biotrend

BioVendor-Laboratorni medicina a.s. CTPark Modrice

BioVision

Bioworld

Bioworlde

BioZol

BMA Biomedicals

Boehringer Mannheim

Bon Opus Biosciences

Boster

Boster

Briar Patch Biosciences

CalBioreagents

Cambio

Cambridge Bioscience

Canada

Cancer Research UK

Capralogics

Caprico Biotechnologies

Cascade Bioscience

Cayman

Cayman Chemical

Cedarlanelabs

Cell Marque

Cell Sciences

Cell Signal

Cell Signaling Technology

Celltechgen

CFTR Folding Consortium

Charles River Laboratories

Chemicon

China

ChromoTek GmbH

Clonegene

Cloud

Cloud-Clone Corp

Cocalico Biologicals

Company

Corning

Cosmo Bio

CovalAb

Creative Biomart

Creativebiolabs

CrownBio

Cruz

Crystal Chem

Cusabio

Cytognos

Cytoskeleton

Cytotech

D-Gen

Dako

DB Biotech

Dbiosys

Dentritics

Detroit R&D

Developmental Studies Hybridoma Bank

Diaclone

Diagenode

Dianova

Diasorin

Diatheva

Eagle BioSciences

Eagle-I

East Coast Biologics

Echelon Biosciences

ECM Biosciences

Eenzyme

EIAab

Eli Lilly

EMD Millipore

Emfret Analytics

EnCor Biotechnology

Enzo Life Sciences

EPC Elastin Products Company

EpiCypher

Epigentek

Equitech-bio

Euro Diagnostica

EuroBioSciences

Eurogentec

Euromedex

Everest Biotech

Evrogen

Exalpha Biologicals

Exbio

Expression Systems

Fabgennix

FibroGen

Fisher

Fitzgerald Industries

Fluidigm

Frontier Institute

Full Moon BioSystems

GE Healthcare Life Biosciences

Gemacbio

Gen-Probe

GeneCopoeia

Genemed

Genentech

GeneTex

Genox

GenScript

Gentaur

Genway Biotech

Genzyme

Germany

Gibco

Gmbh

Haematologic Technologies

Health Protection Agency Culture Collections

Hitachi High Technologies America

Honda

HumanZyme

Hycult Biotech

Hytest

IBL International GmbH

IBT Bioservices

Icosagen

ID Labs

Immundiagnostik

Immune Technology

Immuno-Biological Laboratories

Immunoglobe

ImmunoStar

IMMUNOSTEP

ImmunoTools

Immunovision

Immunoway

ImmunoWay

ImmuQuest

in-house

Inc.

Individual Researcher

Ingenasa

Innogenetics

Innovative Research

Inova Diagnostics

Insightbio

Interchim

International Blood Group Reference Laboratory

Invitrogen

Invitrogen

InvivoGen

J. Sevigny's research lab

Jackson ImmunoResearch Laboratories

Japan

Jena Bioscience

Kamiya Biomedical Company

KeraFAST

Kirkegaard & Perry Laboratories

Kyowa

Laboratories

labs

LAE Biotech

Leica

Leica Biosystems

LI-COR Biosciences

LifeSensors

LifeSpan Biosciences

List Biological Laboratories

Lpbio

Ltd.

Luminex Corporation

Lunginnov

M?diMabs

MABTECH

Mabtechnologies

Maine Biotechnology Services

Matreya

MaxVision Biosciences

MBL International

MD Bioproducts

MD Biosciences

Mediagnost

Medicorp

MedImmune

Menarini

Meridian Life Science

MICROM International GmbH

Millipore

Miltenyi Biotec

Mobitec

Molecular Innovations

Monosan

Moravian Biotechnology

MP Biochemicals

multimmune GmbH

MyBioSource

Nacalai Tesque

Nanoprobes

Nanotools

National Institutes of Health AIDS Research and Reference Reagent Program

Neoclone

Neogeneurope

Neuromab

Neuromics

New England Biolabs

NewEast Biosciences

Nichirei Biosciences Inc.

Nordic BioSite

Nordic-MUbio

Nova Lifetech

Novartis

NovaTec Immundiagnostica GmbH

Novus Biologicals

Nussloch

Ocean Optics Inc.

OriGene

Osenses

Oxisresearch

Panvera

PBL Assay Science

Pel-Freez

PeproTech

PeproTech

PerkinElmer

Perrigo

Perseus Proteomics

Pfizer

Phoenix Pharmaceuticals

Phosphosolutions

Pierce

PIK3CA

Pishtaz Teb Zaman Diagnostics

Polysciences

Precision Antibody

Progen

ProMab

Promega

ProSci

ProSpec

Protein Mods

Protein Sciences

ProteinOne

Proteintech Group

Proteus Biosciences

QED Bioscience

Qiagen

Qiagen

Quartett GmbH

QuickZyme Biosciences

Quidel

R&D Systems

Randox Life Sciences

RayBiotech

RD-Biotech

Recombinant Antibody Network

Reliatech GmbH

ReproCELL

Roche

Roche Applied Science

Rockland Immunochemicals

Sanbio

Sangon

Sanquin

Santa

Santa Cruz Biotechnology

SCETI

SCICONS

Scientific

Scytek

Sekisui Diagnostics (UK) Limited

Selleck

Selleck Chemicals

SICGEN

Sigma

Sigma-Aldrich

SignalChem Pharmaceuticals

Sino Biological

Somru BioScience

Sony

Source BioScience

SouthernBiotech

Spanish National Cancer Research Centre

Speed BioSystems

Spring Bioscience Corp.

St John's Laboratory

Stemcell Technologies

StemRD

Strategic Diagnostics

StressMarq Biosciences

Sungene Biotech

Svar Life Science

SWant

Switzerland

Syd Labs

Symansis

Synaptic Systems

System Biosciences

systems

Taiwan

Takara Bio Clontech

Takara Bio Inc

Tebu

technology

Thermo

Tocris

Tocris Bioscience

Tonbo

Tonbo Biosciences

TopoGEN

Toronto BioScience

Torrey Pines Biolabs

Toxintechnology

TOYOBO

TRANS GENIC

Trendpharmatech

Trevigen

Triple Point Biologics

Tulip Biolabs

US Biological

USA

Vector Laboratories

Ventana

Vincibiochem

Virogen

ViroStat

Virusys

Vision Biosystems

VMRD

Wako Chemicals USA

Wieslab

WILEX Inc.

Wolwobiotech

Wuhan Fine Biotech Co.,Ltd.

Xenotech

Ximbio

YAMASA

YO Proteins

Zebrafish International Resource Center

Zeta Corporation

Zymo Research Corporation

ZYTOMED Systems

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Dictionary