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Article ; Online: A novel assay for drug screening that utilizes the heat shock response of Caenorhabditis elegans nematodes.

Chen, Chih-Hsiung / Patel, Rahul / Bortolami, Alessandro / Sesti, Federico

PloS one

2020 Volume 15, Issue 10, Page(s) e0240255

Abstract: Biological organisms respond to environmental stressors by recruiting multiple cellular cascades that act to mitigate damage and ultimately enhance survival. This implies that compounds that interact with any of those pathways might improve organism's ... ...

Abstract	Biological organisms respond to environmental stressors by recruiting multiple cellular cascades that act to mitigate damage and ultimately enhance survival. This implies that compounds that interact with any of those pathways might improve organism's survival. Here, we report on an initial attempt to develop a drug screening assay based on the heat shock (HS) response of Caenorhabditis elegans nematodes. The protocol works by subjecting the worms to two HS conditions in the absence/presence of the test compounds. Post-heat shock survival is quantified manually or in semi-automatic manner by analyzing z-stack pictures. We blindly screened a cassette of 72 compounds in different developmental stages provided by Eli Lilly through their Open Innovation Drug Discovery program. The analysis indicated that, on average, therapeutically useful drugs increase survival to HS compared to compounds used in non-clinical settings. We developed a formalism that estimates the probability of a compound to enhance survival based on a comparison with a set of parameters calculated from a pool of 35 FDA-approved drugs. The method correctly identified the developmental stages of the Lilly compounds based on their relative abilities to enhance survival to the HS. Taken together these data provide proof of principle that an assay that measures the HS response of C. elegans can offer physiological and pharmacological insight in a cost- and time-efficient manner.
MeSH term(s)	Animals ; Biological Assay/methods ; Caenorhabditis elegans/metabolism ; Caenorhabditis elegans/physiology ; Caenorhabditis elegans Proteins/genetics ; Caenorhabditis elegans Proteins/metabolism ; Drug Evaluation, Preclinical/methods ; Heat-Shock Proteins/genetics ; Heat-Shock Proteins/metabolism ; Heat-Shock Response/genetics ; Heat-Shock Response/physiology
Chemical Substances	Caenorhabditis elegans Proteins ; Heat-Shock Proteins
Language	English
Publishing date	2020-10-09
Publishing country	United States
Document type	Journal Article ; Research Support, N.I.H., Extramural ; Research Support, U.S. Gov't, Non-P.H.S.
ISSN	1932-6203
ISSN (online)	1932-6203
DOI	10.1371/journal.pone.0240255
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: Deletion of ER-retention Motif on SARS-CoV-2 Spike Protein Reduces Cell Hybrid During Cell-cell Fusion.

Chen, Chih-Hsiung / Badeti, Saiaditya / Cho, Jong Hyun / Naghizadeh, Alireza / Wang, Xuening / Liu, Dongfang

Research square

2021

Abstract: The novel SARS-CoV-2 has quickly become a global pandemic since the first reported case in December 2019, with the virus infecting millions of people to date. The spike (S) protein of the SARS-CoV-2 virus plays a key role in binding to angiotensin- ... ...

Abstract	The novel SARS-CoV-2 has quickly become a global pandemic since the first reported case in December 2019, with the virus infecting millions of people to date. The spike (S) protein of the SARS-CoV-2 virus plays a key role in binding to angiotensin-converting enzyme 2 (ACE2), a host cell receptor for SARS-CoV-2. S proteins that are expressed on the cell membrane can initiate receptor-dependent syncytia formation that is associated with extensive tissue damage. Formation of syncytia have been previously observed in cells infected with various other viruses (e.g., HIV, Ebola, Influenza, and Herpesviruses). However, this phenomenon is not well documented and the mechanisms regulating the formation of these syncytia by SARS-CoV-2 are not fully understood. In this study, we investigated the possibility that cell fusion events mediated by the S protein of SARS-CoV-2 and ACE2 interaction can occur in different human cell lines that mimic different tissue origins. These cell lines were stably transduced with either wild-type (WT-S) S protein or a mutated variant where the ER-retention motif was removed (Δ19-S), or human ACE2 vectors. Different co-culture combinations of spike-expressing 293T, A549, K562, and SK-Hep1 cells with hACE2-expressing cells revealed cell hybrid fusion. However, only certain cells expressing S protein can form syncytial structures as this phenomenon cannot be observed in all co-culture combinations. Thus, SARS-CoV-2 mediated cell-cell fusion represents a cell type-dependent process which might rely on a different set of parameters. Recently, the Δ19-S variant is being widely used to increase SARS-CoV-2 pseudovirus production for in vitro assays. Comparison of cell fusion occurring via Δ19-S expressing cells shows defective nuclear fusion and syncytia formation compared to WT-S. This distinction between the Δ19-S variant and WT-S protein may have downstream implications for studies that utilize pseudovirus-based entry assays. Additionally, this study suggest that spike protein expressed by vaccines may affect different ACE2-expressing host cells after SARS-CoV-2 vaccine administration. The long-term effects of these vaccines should be monitored carefully.
Language	English
Publishing date	2021-04-09
Publishing country	United States
Document type	Preprint
DOI	10.21203/rs.3.rs-380389/v1
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: S309-CAR-NK cells bind the Omicron variants

Ma, Minh Tuyet / Jiang, Qingkui / Chen, Chih-Hsiung / Badeti, Saiaditya / Wang, Xuening / Zeng, Cong / Evans, Deborah / Bodnar, Brittany / Marras, Salvatore A E / Tyagi, Sanjay / Bharaj, Preeti / Yehia, Ghassan / Romanienko, Peter / Hu, Wenhui / Liu, Shan-Lu / Shi, Lanbo / Liu, Dongfang

Journal of virology

2024 , Page(s) e0003824

Abstract: Recent progress on chimeric antigen receptor (CAR)-NK cells has shown promising results in treating CD19-positive lymphoid tumors with minimal toxicities [including graft versus host disease (GvHD) and cytokine release syndrome (CRS) in clinical trials. ... ...

Abstract	Recent progress on chimeric antigen receptor (CAR)-NK cells has shown promising results in treating CD19-positive lymphoid tumors with minimal toxicities [including graft versus host disease (GvHD) and cytokine release syndrome (CRS) in clinical trials. Nevertheless, the use of CAR-NK cells in combating viral infections has not yet been fully explored. Previous studies have shown that CAR-NK cells expressing S309 single-chain fragment variable (scFv), hereinafter S309-CAR-NK cells, can bind to SARS-CoV-2 wildtype pseudotyped virus (PV) and effectively kill cells expressing wild-type spike protein Importance: Chimeric antigen receptor (CAR)-NK cells can be "off-the-shelf" products that treat various diseases, including cancer, infections, and autoimmune diseases. In this study, we engineered natural killer (NK) cells to express S309 single-chain fragment variable (scFv), to target the Spike protein of SARS-CoV-2, hereinafter S309-CAR-NK cells. Our study shows that S309-CAR-NK cells are effective against different SARS-CoV-2 variants, including the B.1.617.2 (Delta), B.1.621 (Mu), and B.1.1.529 (Omicron) variants. The S309-CAR-NK cells can (i) directly bind to SARS-CoV-2 pseudotyped virus (PV), (ii) competitively bind to SARS-CoV-2 PV with 293T cells expressing the human angiotensin-converting enzyme 2 (hACE2) receptor (293T-hACE2 cells), (iii) specifically target and lyse A549 cells expressing the spike protein, and (iv) significantly reduce the viral loads of SARS-CoV-2 wild-type (strain USA/WA1/2020) in the lungs of NOD/SCID gamma (NSG) mice expressing hACE2 (hACE2-NSG mice). Altogether, the current study demonstrates the potential use of S309-CAR-NK immunotherapy as an alternative treatment for COVID-19 patients.
Language	English
Publishing date	2024-05-20
Publishing country	United States
Document type	Journal Article
ZDB-ID	80174-4
ISSN	1098-5514 ; 0022-538X
ISSN (online)	1098-5514
ISSN	0022-538X
DOI	10.1128/jvi.00038-24
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Article: Phylogeny and Historical Biogeography of

Tsai, Chi-Chu / Liao, Pei-Chun / Ko, Ya-Zhu / Chen, Chih-Hsiung / Chiang, Yu-Chung

Frontiers in plant science

2020 Volume 11, Page(s) 126

Abstract: The phylogeny and biogeography of the ... ...

Abstract	The phylogeny and biogeography of the genus
Language	English
Publishing date	2020-02-27
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	2711035-7
ISSN	1664-462X
ISSN	1664-462X
DOI	10.3389/fpls.2020.00126
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Article ; Online: Modeling of ACE2 and antibodies bound to SARS-CoV-2 provides insights into infectivity and immune evasion.

Lubin, Joseph H / Markosian, Christopher / Balamurugan, D / Ma, Minh T / Chen, Chih-Hsiung / Liu, Dongfang / Pasqualini, Renata / Arap, Wadih / Burley, Stephen K / Khare, Sagar D

JCI insight

2023 Volume 8, Issue 13

Abstract: Given the COVID-19 pandemic, there is interest in understanding ligand-receptor features and targeted antibody-binding attributes against emerging SARS-CoV-2 variants. Here, we developed a large-scale structure-based pipeline for analysis of protein- ... ...

Abstract	Given the COVID-19 pandemic, there is interest in understanding ligand-receptor features and targeted antibody-binding attributes against emerging SARS-CoV-2 variants. Here, we developed a large-scale structure-based pipeline for analysis of protein-protein interactions regulating SARS-CoV-2 immune evasion. First, we generated computed structural models of the Spike protein of 3 SARS-CoV-2 variants (B.1.1.529, BA.2.12.1, and BA.5) bound either to a native receptor (ACE2) or to a large panel of targeted ligands (n = 282), which included neutralizing or therapeutic monoclonal antibodies. Moreover, by using the Barnes classification, we noted an overall loss of interfacial interactions (with gain of new interactions in certain cases) at the receptor-binding domain (RBD) mediated by substituted residues for neutralizing complexes in classes 1 and 2, whereas less destabilization was observed for classes 3 and 4. Finally, an experimental validation of predicted weakened therapeutic antibody binding was performed in a cell-based assay. Compared with the original Omicron variant (B.1.1.529), derivative variants featured progressive destabilization of antibody-RBD interfaces mediated by a larger set of substituted residues, thereby providing a molecular basis for immune evasion. This approach and findings provide a framework for rapidly and efficiently generating structural models for SARS-CoV-2 variants bound to ligands of mechanistic and therapeutic value.
MeSH term(s)	Humans ; SARS-CoV-2 ; Angiotensin-Converting Enzyme 2 ; COVID-19 ; Immune Evasion ; Ligands ; Pandemics ; Antibodies, Monoclonal
Chemical Substances	Angiotensin-Converting Enzyme 2 (EC 3.4.17.23) ; Ligands ; Antibodies, Monoclonal
Language	English
Publishing date	2023-07-10
Publishing country	United States
Document type	Journal Article ; Research Support, U.S. Gov't, Non-P.H.S. ; Research Support, N.I.H., Extramural ; Research Support, Non-U.S. Gov't
ISSN	2379-3708
ISSN (online)	2379-3708
DOI	10.1172/jci.insight.168296
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Article: Deletion of ER-retention motif on SARS-CoV-2 spike protein reduces cell hybrid during cell-cell fusion.

Wang, Xuening / Chen, Chih-Hsiung / Badeti, Saiaditya / Cho, Jong Hyun / Naghizadeh, Alireza / Wang, Ziren / Liu, Dongfang

Cell & bioscience

2021 Volume 11, Issue 1, Page(s) 114

Abstract: Background: The novel SARS-CoV-2 has quickly become a global pandemic since the first reported case in December 2019, with the virus infecting millions of people to date. The spike (S) protein of the SARS-CoV-2 virus plays a key role in binding to ... ...

Abstract	Background: The novel SARS-CoV-2 has quickly become a global pandemic since the first reported case in December 2019, with the virus infecting millions of people to date. The spike (S) protein of the SARS-CoV-2 virus plays a key role in binding to angiotensin-converting enzyme 2 (ACE2), a host cell receptor for SARS-CoV-2. S proteins that are expressed on the cell membrane can initiate receptor-dependent syncytia formation that is associated with extensive tissue damage. Formation of syncytia have been previously observed in cells infected with various other viruses (e.g., HIV, Ebola, Influenza, and Herpesviruses). However, this phenomenon is not well documented and the mechanisms regulating the formation of the syncytia by SARS-CoV-2 are not fully understood. Results: In this study, we investigated the possibility that cell fusion events mediated by the S protein of SARS-CoV-2 and ACE2 interaction can occur in different human cell lines that mimic different tissue origins. These cell lines were transduced with either wild-type (WT-S) S protein or a mutated variant where the ER-retention motif was removed (Δ19-S), as well as human ACE2 expression vectors. Different co-culture combinations of spike-expressing 293T, A549, K562, and SK-Hep1 cells with hACE2-expressing cells revealed cell hybrid fusion. However, only certain cells expressing S protein can form syncytial structures as this phenomenon cannot be observed in all co-culture combinations. Thus, SARS-CoV-2 mediated cell-cell fusion represents a cell type-dependent process which might rely on a different set of parameters. Recently, the Δ19-S variant is being widely used to increase SARS-CoV-2 pseudovirus production for in vitro assays. Comparison of cell fusion occurring via Δ19-S expressing cells shows defective nuclear fusion and syncytia formation compared to WT-S. Conclusions: This distinction between the Δ19-S variant and WT-S protein may have downstream implications for studies that utilize pseudovirus-based entry assays. Additionally, this study suggest that spike protein expressed by vaccines may affect different ACE2-expressing host cells after SARS-CoV-2 vaccine administration. The long-term effects of these vaccines should be monitored carefully. Δ19-S mRNA may represent a safer mRNA vaccine design in the future.
Language	English
Publishing date	2021-06-23
Publishing country	England
Document type	Journal Article
ZDB-ID	2593367-X
ISSN	2045-3701
ISSN	2045-3701
DOI	10.1186/s13578-021-00626-0
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Article: CAR-NK Cells Effectively Target the D614 and G614 SARS-CoV-2-infected Cells.

Ma, Minh / Badeti, Saiaditya / Chen, Chih-Hsiung / Pinter, Abraham / Jiang, Qingkui / Shi, Lanbo / Zhou, Renping / Xu, Huanbin / Li, Qingsheng / Gause, William / Liu, Dongfang

bioRxiv : the preprint server for biology

2021

Abstract: Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is highly contagious presenting a significant public health issue. Current therapies used to treat coronavirus disease 2019 (COVID-19) include monoclonal antibody cocktail, convalescent plasma, ...

Abstract	Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is highly contagious presenting a significant public health issue. Current therapies used to treat coronavirus disease 2019 (COVID-19) include monoclonal antibody cocktail, convalescent plasma, antivirals, immunomodulators, and anticoagulants, though the current therapeutic options remain limited and expensive. The vaccines from Pfizer and Moderna have recently been authorized for emergency use, which are invaluable for the prevention of SARS-CoV-2 infection. However, their long-term side effects are not yet to be documented, and populations with immunocompromised conditions (e.g., organ-transplantation and immunodeficient patients) may not be able to mount an effective immune response. In addition, there are concerns that wide-scale immunity to SARS-CoV-2 may introduce immune pressure that could select for escape mutants to the existing vaccines and monoclonal antibody therapies. Emerging evidence has shown that chimeric antigen receptor (CAR)- natural killer (NK) immunotherapy has potent antitumor response in hematologic cancers with minimal adverse effects in recent studies, however, the potentials of CAR-NK cells in preventing and treating severe cases of COVID-19 has not yet been fully exploited. Here, we improve upon a novel approach for the generation of CAR-NK cells for targeting SARS-CoV-2 and its D614G mutant. CAR-NK cells were generated using the scFv domain of S309 (henceforward, S309-CAR-NK), a SARS-CoV and SARS-CoV-2 neutralizing antibody that targets the highly conserved region of SARS-CoV-2 spike (S) glycoprotein, therefore would be more likely to recognize different variants of SARS-CoV-2 isolates. S309-CAR-NK cells can specifically bind to pseudotyped SARS-CoV-2 virus and its D614G mutant. Furthermore, S309-CAR-NK cells can specifically kill target cells expressing SARS-CoV-2 S protein
Language	English
Publishing date	2021-01-15
Publishing country	United States
Document type	Preprint
DOI	10.1101/2021.01.14.426742
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: CAR-NK Cells Effectively Target SARS-CoV-2-Spike-Expressing Cell Lines

Ma, Minh Tuyet / Badeti, Saiaditya / Chen, Chih-Hsiung / Kim, James / Choudhary, Alok / Honnen, Bill / Reichman, Charles / Calianese, David / Pinter, Abraham / Jiang, Qingkui / Shi, Lanbo / Zhou, Renping / Xu, Huanbin / Li, Qingsheng / Gause, William / Liu, Dongfang

Frontiers in immunology

2021 Volume 12, Page(s) 652223

Abstract: Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is highly contagious and presents a significant public health issue. Current therapies used to treat coronavirus disease 2019 (COVID-19) include monoclonal antibody cocktail, convalescent ... ...

Abstract	Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is highly contagious and presents a significant public health issue. Current therapies used to treat coronavirus disease 2019 (COVID-19) include monoclonal antibody cocktail, convalescent plasma, antivirals, immunomodulators, and anticoagulants. The vaccines from Pfizer and Moderna have recently been authorized for emergency use, which are invaluable for the prevention of SARS-CoV-2 infection. However, their long-term side effects are not yet documented, and populations with immunocompromised conditions (e.g., organ-transplantation and immunodeficient patients) may not be able to mount an effective immune response. In addition, there are concerns that wide-scale immunity to SARS-CoV-2 may introduce immune pressure that could select for escape mutants to the existing vaccines and monoclonal antibody therapies. Emerging evidence has shown that chimeric antigen receptor (CAR)- natural killer (NK) immunotherapy has potent antitumor response in hematologic cancers with minimal adverse effects in recent studies, however, the potentials of CAR-NK cells in treating COVID-19 has not yet been fully exploited. Here, we improve upon a novel approach for the generation of CAR-NK cells for targeting SARS-CoV-2 and its various mutants. CAR-NK cells were generated using the scFv domain of S309 (henceforward, S309-CAR-NK), a SARS-CoV and SARS-CoV-2 neutralizing antibody (NAbs) that targets the highly conserved region of SARS-CoV-2 spike (S) glycoprotein and is therefore more likely to recognize different variants of SARS-CoV-2 isolates. S309-CAR-NK cells can specifically bind to pseudotyped SARS-CoV-2 virus and its D614G, N501Y, and E484K mutants. Furthermore, S309-CAR-NK cells can specifically kill target cells expressing SARS-CoV-2 S protein
MeSH term(s)	A549 Cells ; COVID-19/genetics ; COVID-19/immunology ; COVID-19/pathology ; COVID-19/therapy ; Gene Expression Regulation/genetics ; Gene Expression Regulation/immunology ; Hep G2 Cells ; Humans ; Killer Cells, Natural/immunology ; Receptors, Chimeric Antigen/genetics ; Receptors, Chimeric Antigen/immunology ; SARS-CoV-2/genetics ; SARS-CoV-2/immunology ; Spike Glycoprotein, Coronavirus/genetics ; Spike Glycoprotein, Coronavirus/immunology
Chemical Substances	Receptors, Chimeric Antigen ; Spike Glycoprotein, Coronavirus ; spike protein, SARS-CoV-2
Language	English
Publishing date	2021-07-23
Publishing country	Switzerland
Document type	Journal Article ; Research Support, N.I.H., Extramural ; Research Support, Non-U.S. Gov't
ZDB-ID	2606827-8
ISSN	1664-3224 ; 1664-3224
ISSN (online)	1664-3224
ISSN	1664-3224
DOI	10.3389/fimmu.2021.652223
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: CAR-NK Cells Effectively Target the D614 and G614 SARS-CoV-2-infected Cells

Ma, Minh / Badeti, Saiaditya / Chen, Chih-Hsiung / Pinter, Abraham / Jiang, Qingkui / Shi, Lanbo / Zhou, Renping / Xu, Huanbin / Li, Qingsheng / Gause, William / Liu, Dongfang

bioRxiv

Abstract	Severe Acute Respiratory Syndrome Coronavirus 2 (SARS-CoV-2) is highly contagious presenting a significant public health issue. Current therapies used to treat coronavirus disease 2019 (COVID-19) include monoclonal antibody cocktail, convalescent plasma, antivirals, immunomodulators, and anticoagulants, though the current therapeutic options remain limited and expensive. The vaccines from Pfizer and Moderna have recently been authorized for emergency use, which are invaluable for the prevention of SARS-CoV-2 infection. However, their long-term side effects are not yet to be documented, and populations with immunocompromised conditions (e.g., organ-transplantation and immunodeficient patients) may not be able to mount an effective immune response. In addition, there are concerns that wide-scale immunity to SARS-CoV-2 may introduce immune pressure that could select for escape mutants to the existing vaccines and monoclonal antibody therapies. Emerging evidence has shown that chimeric antigen receptor (CAR)- natural killer (NK) immunotherapy has potent antitumor response in hematologic cancers with minimal adverse effects in recent studies, however, the potentials of CAR-NK cells in preventing and treating severe cases of COVID-19 has not yet been fully exploited. Here, we improve upon a novel approach for the generation of CAR-NK cells for targeting SARS-CoV-2 and its D614G mutant. CAR-NK cells were generated using the scFv domain of S309 (henceforward, S309-CAR-NK), a SARS-CoV and SARS-CoV-2 neutralizing antibody that targets the highly conserved region of SARS-CoV-2 spike (S) glycoprotein, therefore would be more likely to recognize different variants of SARS-CoV-2 isolates. S309-CAR-NK cells can specifically bind to pseudotyped SARS-CoV-2 virus and its D614G mutant. Furthermore, S309-CAR-NK cells can specifically kill target cells expressing SARS-CoV-2 S protein in vitro and show superior killing activity and cytokine production, compared to that of the recently published CR3022-CAR-NK cells. Thus, these results pave the way for generating off-the-shelf S309-CAR-NK cells for treatment in high-risk individuals as well as provide an alternative strategy for patients unresponsive to current vaccines.
Keywords	covid19
Language	English
Publishing date	2021-01-15
Publisher	Cold Spring Harbor Laboratory
Document type	Article ; Online
DOI	10.1101/2021.01.14.426742
Database	COVID19

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Article: Out‐of‐Tibet: the spatio‐temporal evolution of Gentiana (Gentianaceae)

Favre, Adrien / Chen, Chih‐Hsiung / Matuszak, Sabine / Michalak, Ingo / Muellner‐Riehl, Alexandra N / Pringle, James S / Struwe, Lena / Sun, Hang / Wang, Jenn‐Che / Yuan, Yong‐Ming

Journal of biogeography. 2016 Oct., v. 43, no. 10

2016

Abstract: AIM: We investigated the historical biogeography and diversification of Gentiana L. (Gentianaceae). Our study depicts the origin and dispersal routes of this alpine genus, and the role of the uplift of the Qinghai–Tibet Plateau (QTP) and past climate ... ...

Abstract	AIM: We investigated the historical biogeography and diversification of Gentiana L. (Gentianaceae). Our study depicts the origin and dispersal routes of this alpine genus, and the role of the uplift of the Qinghai–Tibet Plateau (QTP) and past climate changes as triggers for its diversification. LOCATION: Tibeto‐Himalayan region and world‐wide mountain habitats. METHODS: Our sampling represents more than 50% of the extant Gentiana species, including all sections across their entire geographical ranges. We investigated the evolutionary history of Gentiana using phylogenetic reconstructions (maximum likelihood and Bayesian inference) of ITS, atpB–rbcL and trnL–trnF sequences, as well as molecular dating with beast. We tested two approaches of ancestral area reconstructions (DEC, DIVA) in BioGeoBEARS and investigated diversification rates using BAMM. RESULTS: The common ancestor of Gentiana and subtribe Gentianinae lived in the QTP region at around 34 (25–45) million years ago (Ma), and 40 (29–52) Ma respectively. From the surroundings of the QTP, Gentiana lineages dispersed to eastern China, Taiwan, Europe, North and South America, Australia and New Guinea, from mid‐Miocene onward (c. 15 Ma–present), with only one older dispersal event to Europe (c. 37–21 Ma). Diversification rates gradually increased over time, and two switches of diversification rates were identified in Gentianinae (c. 7 Ma, simultaneously in the Pneumonanthe/Cruciata lineage and in Tripterospermum). MAIN CONCLUSIONS: Gentiana existed in the QTP region throughout most of its uplift history following the India‐Asia collision. This region acted as the primary source area for dispersals to many areas of the world. Because steady increase in diversification rates coincides with the extension of the QTP, we argue that the museum theory rather than the explosive radiation theory prevails for gentians in this region, although rare shifts of diversification rates are associated with niche shifts across the alpine/subalpine ecotone.
Keywords	Bayesian theory ; biogeography ; climate change ; ecotones ; Gentiana ; habitats ; internal transcribed spacers ; phylogeny ; Australia ; China ; Europe ; New Guinea ; South America ; Taiwan
Language	English
Dates of publication	2016-10
Size	p. 1967-1978.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	188963-1
ISSN	0305-0270
ISSN	0305-0270
DOI	10.1111/jbi.12840
Database	NAL-Catalogue (AGRICOLA)

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