Developmental bias in the evolution and plasticity of beetle horn shape
The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject to much debate. Here, we assess whether developmental variability in beetle horn shape aligns with the phenotypic effects of plasticity and...
Saved in:
Published in | Proceedings of the Royal Society. B, Biological sciences Vol. 289; no. 1983; p. 20221441 |
---|---|
Main Authors | , , |
Format | Journal Article |
Language | English |
Published |
England
The Royal Society
28.09.2022
|
Subjects | |
Online Access | Get full text |
ISSN | 0962-8452 1471-2954 1471-2954 |
DOI | 10.1098/rspb.2022.1441 |
Cover
Loading…
Abstract | The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject to much debate. Here, we assess whether developmental variability in beetle horn shape aligns with the phenotypic effects of plasticity and evolutionary divergence, yielding three salient results. First, we find that most pathways previously shown to regulate horn length also affect shape. Second, we find that the phenotypic effects of manipulating divergent developmental pathways are correlated with each other as well as multivariate fluctuating asymmetry—a measure of developmental variability. Third, these effects further aligned with thermal plasticity, population differences and macroevolutionary divergence between sister taxa and more distantly related species. Collectively, our results support the hypothesis that changes in horn shape—whether brought about by environmentally plastic responses, functional manipulations or evolutionary divergences—converge along ‘developmental lines of least resistance’, i.e. are biased by the developmental system underpinning horn shape. |
---|---|
AbstractList | The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject to much debate. Here, we assess whether developmental variability in beetle horn shape aligns with the phenotypic effects of plasticity and evolutionary divergence, yielding three salient results. First, we find that most pathways previously shown to regulate horn length also affect shape. Second, we find that the phenotypic effects of manipulating divergent developmental pathways are correlated with each other as well as multivariate fluctuating asymmetry-a measure of developmental variability. Third, these effects further aligned with thermal plasticity, population differences and macroevolutionary divergence between sister taxa and more distantly related species. Collectively, our results support the hypothesis that changes in horn shape-whether brought about by environmentally plastic responses, functional manipulations or evolutionary divergences-converge along 'developmental lines of least resistance', i.e. are biased by the developmental system underpinning horn shape.The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject to much debate. Here, we assess whether developmental variability in beetle horn shape aligns with the phenotypic effects of plasticity and evolutionary divergence, yielding three salient results. First, we find that most pathways previously shown to regulate horn length also affect shape. Second, we find that the phenotypic effects of manipulating divergent developmental pathways are correlated with each other as well as multivariate fluctuating asymmetry-a measure of developmental variability. Third, these effects further aligned with thermal plasticity, population differences and macroevolutionary divergence between sister taxa and more distantly related species. Collectively, our results support the hypothesis that changes in horn shape-whether brought about by environmentally plastic responses, functional manipulations or evolutionary divergences-converge along 'developmental lines of least resistance', i.e. are biased by the developmental system underpinning horn shape. The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject to much debate. Here, we assess whether developmental variability in beetle horn shape aligns with the phenotypic effects of plasticity and evolutionary divergence, yielding three salient results. First, we find that most pathways previously shown to regulate horn length also affect shape. Second, we find that the phenotypic effects of manipulating divergent developmental pathways are correlated with each other as well as multivariate fluctuating asymmetry—a measure of developmental variability. Third, these effects further aligned with thermal plasticity, population differences and macroevolutionary divergence between sister taxa and more distantly related species. Collectively, our results support the hypothesis that changes in horn shape—whether brought about by environmentally plastic responses, functional manipulations or evolutionary divergences—converge along ‘developmental lines of least resistance’, i.e. are biased by the developmental system underpinning horn shape. |
Author | Moczek, Armin P. Rohner, Patrick T. Hu, Yonggang |
Author_xml | – sequence: 1 givenname: Patrick T. orcidid: 0000-0002-9840-1050 surname: Rohner fullname: Rohner, Patrick T. organization: Department of Biology, Indiana University Bloomington, Bloomington, IN 47405, USA – sequence: 2 givenname: Yonggang orcidid: 0000-0002-3438-7296 surname: Hu fullname: Hu, Yonggang organization: Department of Biology, Indiana University Bloomington, Bloomington, IN 47405, USA, State Key Laboratory of Silkworm Genome Biology, Institute of Sericulture and Systems Biology, Southwest University, Chongqing 400715, People's Republic of China – sequence: 3 givenname: Armin P. orcidid: 0000-0002-3478-9949 surname: Moczek fullname: Moczek, Armin P. organization: Department of Biology, Indiana University Bloomington, Bloomington, IN 47405, USA |
BackLink | https://www.ncbi.nlm.nih.gov/pubmed/36168764$$D View this record in MEDLINE/PubMed |
BookMark | eNp1kUlLxDAYhoMoOi5Xj5Kjl45Zm-QiiDsIXvQc0varE8kktekMzL-3xQUVPOXwvUt43n20HVMEhI4pmVNi9Fmfu2rOCGNzKgTdQjMqFC2YkWIbzYgpWaGFZHtoP-dXQoiRWu6iPV7SUqtSzNDtFawhpG4JcXABV95l7CMeFoBhncJq8CliFxvcBZcHX_thg1OLK4AhAF6kPuK8cB0cop3WhQxHn-8Ber65frq8Kx4eb-8vLx6KWnAxFI2TmrhWOEZaaiQQRSrRiFIazWtueAVcSK14WzeKEC0Z4bqWXEnWtpVSgh-g84_cblUtoanHb_cu2K73S9dvbHLe_r5Ev7AvaW2NpLLkZAw4_Qzo09sK8mCXPtcQgouQVtkyRfWIjZVmlJ787Pou-aI3CsSHoO5Tzj20duTjJmRjtQ-WEjuNZKeR7DSSnUYabfM_tq_kfwzv_BWUuQ |
CitedBy_id | crossref_primary_10_2992_007_090_0201 crossref_primary_10_3389_fcell_2024_1453566 crossref_primary_10_1016_j_cois_2023_101114 crossref_primary_10_1098_rspb_2022_1441 crossref_primary_10_1186_s13227_024_00223_5 crossref_primary_10_1038_s41559_025_02639_1 crossref_primary_10_1111_ede_12436 crossref_primary_10_1111_pala_12719 crossref_primary_10_1111_ede_12464 |
Cites_doi | 10.1111/j.1558-5646.1998.tb02018.x 10.1111/j.1525-142X.2007.00176.x 10.1111/j.1420-9101.2008.01573.x 10.1673/2006_06_21.1 10.1098/rstb.2013.0249 10.1111/evo.14147 10.3389/fevo.2019.00056 10.1016/j.cois.2016.11.004 10.1111/j.1558-5646.2007.00071.x 10.1093/oso/9780198548959.001.0001 10.1126/science.1056607 10.1111/j.1558-5646.1979.tb04694.x 10.1073/pnas.1315213110 10.1111/j.1558-5646.2012.01838.x 10.1073/pnas.1403667112 10.1007/s00427-016-0539-2 10.3390/sym7020843 10.1073/pnas.2008253117 10.1093/biolinnean/blaa228 10.1074/jbc.M117.777797 10.1371/journal.pbio.1000365 10.1007/s00427-020-00645-w 10.1073/pnas.1409585111 10.1016/S0022-5193(84)80050-8 10.1098/rspb.2021.0241 10.2307/2410734 10.1006/anbe.1999.1342 10.1098/rspb.2022.1441 10.1534/genetics.118.300995 10.1111/jeb.13429 10.1073/pnas.0809668106 10.1111/j.0014-3820.2005.tb01044.x 10.5962/p.372750 10.1111/j.0014-3820.2005.tb01763.x 10.1038/nature23473 10.1016/j.ympev.2016.08.016 10.1016/j.tig.2012.02.002 10.1146/annurev.med.60.042307.110802 10.1093/sysbio/syt025 10.1086/386551 10.1098/rspb.2015.1119 10.1002/jez.2413 10.1098/rspb.2018.1631 10.1111/j.1095-8312.2006.00674.x 10.1111/evo.14045 10.1086/414425 10.1073/pnas.1118589109 10.1038/ncomms4700 |
ContentType | Journal Article |
Copyright | 2022 The Author(s) 2022 |
Copyright_xml | – notice: 2022 The Author(s) 2022 |
DBID | AAYXX CITATION CGR CUY CVF ECM EIF NPM 7X8 5PM |
DOI | 10.1098/rspb.2022.1441 |
DatabaseName | CrossRef Medline MEDLINE MEDLINE (Ovid) MEDLINE MEDLINE PubMed MEDLINE - Academic PubMed Central (Full Participant titles) |
DatabaseTitle | CrossRef MEDLINE Medline Complete MEDLINE with Full Text PubMed MEDLINE (Ovid) MEDLINE - Academic |
DatabaseTitleList | MEDLINE - Academic CrossRef MEDLINE |
Database_xml | – sequence: 1 dbid: NPM name: PubMed url: https://proxy.k.utb.cz/login?url=http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed sourceTypes: Index Database – sequence: 2 dbid: EIF name: MEDLINE url: https://proxy.k.utb.cz/login?url=https://www.webofscience.com/wos/medline/basic-search sourceTypes: Index Database |
DeliveryMethod | fulltext_linktorsrc |
Discipline | Sciences (General) Biology |
DocumentTitleAlternate | Developmental bias in the evolution and plasticity of beetle horn shape |
EISSN | 1471-2954 |
ExternalDocumentID | PMC9515630 36168764 10_1098_rspb_2022_1441 |
Genre | Research Support, U.S. Gov't, Non-P.H.S Research Support, Non-U.S. Gov't Journal Article |
GrantInformation_xml | – fundername: ; grantid: P400PB_199257 – fundername: ; grantid: 61369 – fundername: ; grantid: 1901680; IOS 1256689 |
GroupedDBID | --- -~X 0R~ 29P 2WC 36Y 4.4 5RE 85S AACGO AANCE AAYXX ABPLY ABTLG ACIWK ACNCT ACPRK ACQIA ADBBV ADIYS AFRAH ALMA_UNASSIGNED_HOLDINGS ALMYZ AOIJS BAWUL BTFSW CITATION CS3 DIK E3Z EBS F5P FRP GX1 H13 HYE HZ~ JLS JSG KQ8 MRS O9- OK1 RPM TR2 W8F ~02 CGR CUY CVF ECM EIF NPM 7X8 5PM |
ID | FETCH-LOGICAL-c434t-da580af4a20f195e070b4d465983c393be345873fcd700852038c53752ffb7743 |
ISSN | 0962-8452 1471-2954 |
IngestDate | Thu Aug 21 18:36:31 EDT 2025 Thu Jul 10 22:13:21 EDT 2025 Mon Jul 21 06:02:11 EDT 2025 Thu Apr 24 22:59:04 EDT 2025 Tue Jul 01 01:05:21 EDT 2025 |
IsDoiOpenAccess | false |
IsOpenAccess | true |
IsPeerReviewed | true |
IsScholarly | true |
Issue | 1983 |
Keywords | integration developmental bias fluctuating asymmetry geometric morphometrics developmental plasticity Onthophagus |
Language | English |
License | Published by the Royal Society. All rights reserved. |
LinkModel | OpenURL |
MergedId | FETCHMERGED-LOGICAL-c434t-da580af4a20f195e070b4d465983c393be345873fcd700852038c53752ffb7743 |
Notes | ObjectType-Article-1 SourceType-Scholarly Journals-1 ObjectType-Feature-2 content type line 23 Electronic supplementary material is available online at https://doi.org/10.6084/m9.figshare.c.6210975. |
ORCID | 0000-0002-9840-1050 0000-0002-3438-7296 0000-0002-3478-9949 |
OpenAccessLink | https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2022.1441 |
PMID | 36168764 |
PQID | 2718962269 |
PQPubID | 23479 |
ParticipantIDs | pubmedcentral_primary_oai_pubmedcentral_nih_gov_9515630 proquest_miscellaneous_2718962269 pubmed_primary_36168764 crossref_citationtrail_10_1098_rspb_2022_1441 crossref_primary_10_1098_rspb_2022_1441 |
ProviderPackageCode | CITATION AAYXX |
PublicationCentury | 2000 |
PublicationDate | 2022-09-28 20220928 |
PublicationDateYYYYMMDD | 2022-09-28 |
PublicationDate_xml | – month: 09 year: 2022 text: 2022-09-28 day: 28 |
PublicationDecade | 2020 |
PublicationPlace | England |
PublicationPlace_xml | – name: England |
PublicationTitle | Proceedings of the Royal Society. B, Biological sciences |
PublicationTitleAlternate | Proc Biol Sci |
PublicationYear | 2022 |
Publisher | The Royal Society |
Publisher_xml | – name: The Royal Society |
References | e_1_3_6_30_2 e_1_3_6_51_2 e_1_3_6_53_2 Tyndale-Biscoe M (e_1_3_6_50_2) 1996 Qiu W (e_1_3_6_40_2) 2020 e_1_3_6_19_2 e_1_3_6_13_2 e_1_3_6_38_2 e_1_3_6_59_2 e_1_3_6_17_2 e_1_3_6_55_2 e_1_3_6_36_2 e_1_3_6_57_2 e_1_3_6_21_2 e_1_3_6_42_2 e_1_3_6_4_2 e_1_3_6_2_2 e_1_3_6_8_2 e_1_3_6_6_2 e_1_3_6_27_2 e_1_3_6_48_2 Camara MD (e_1_3_6_11_2) 1999; 53 e_1_3_6_29_2 Claude J (e_1_3_6_35_2) 2008 e_1_3_6_23_2 e_1_3_6_44_2 e_1_3_6_25_2 e_1_3_6_46_2 e_1_3_6_52_2 e_1_3_6_54_2 e_1_3_6_10_2 Collyer ML (e_1_3_6_34_2) 2019 Rohlf FJ (e_1_3_6_31_2) 2009 Fincher G (e_1_3_6_49_2) 1975; 29 Moczek AP (e_1_3_6_43_2) 2009 e_1_3_6_14_2 e_1_3_6_37_2 e_1_3_6_12_2 e_1_3_6_39_2 e_1_3_6_18_2 e_1_3_6_33_2 e_1_3_6_56_2 e_1_3_6_16_2 e_1_3_6_58_2 e_1_3_6_41_2 e_1_3_6_20_2 Adams DC (e_1_3_6_32_2) 2021 e_1_3_6_5_2 e_1_3_6_3_2 e_1_3_6_9_2 e_1_3_6_7_2 Møller AP (e_1_3_6_15_2) 1997 e_1_3_6_26_2 e_1_3_6_28_2 e_1_3_6_22_2 e_1_3_6_45_2 e_1_3_6_24_2 e_1_3_6_47_2 |
References_xml | – ident: e_1_3_6_16_2 doi: 10.1111/j.1558-5646.1998.tb02018.x – volume-title: RRPP: linear model evaluation with randomized residuals in a permutation procedure year: 2019 ident: e_1_3_6_34_2 – ident: e_1_3_6_5_2 doi: 10.1111/j.1525-142X.2007.00176.x – ident: e_1_3_6_39_2 doi: 10.1111/j.1420-9101.2008.01573.x – ident: e_1_3_6_56_2 doi: 10.1673/2006_06_21.1 – ident: e_1_3_6_20_2 doi: 10.1098/rstb.2013.0249 – ident: e_1_3_6_38_2 doi: 10.1111/evo.14147 – ident: e_1_3_6_14_2 doi: 10.3389/fevo.2019.00056 – ident: e_1_3_6_22_2 doi: 10.1016/j.cois.2016.11.004 – ident: e_1_3_6_13_2 doi: 10.1111/j.1558-5646.2007.00071.x – volume-title: Asymmetry, developmental stability and evolution year: 1997 ident: e_1_3_6_15_2 doi: 10.1093/oso/9780198548959.001.0001 – ident: e_1_3_6_55_2 doi: 10.1126/science.1056607 – ident: e_1_3_6_8_2 doi: 10.1111/j.1558-5646.1979.tb04694.x – ident: e_1_3_6_3_2 doi: 10.1073/pnas.1315213110 – ident: e_1_3_6_12_2 doi: 10.1111/j.1558-5646.2012.01838.x – ident: e_1_3_6_17_2 doi: 10.1073/pnas.1403667112 – ident: e_1_3_6_33_2 doi: 10.1007/s00427-016-0539-2 – ident: e_1_3_6_19_2 doi: 10.3390/sym7020843 – ident: e_1_3_6_6_2 doi: 10.1073/pnas.2008253117 – ident: e_1_3_6_42_2 doi: 10.1093/biolinnean/blaa228 – ident: e_1_3_6_45_2 doi: 10.1074/jbc.M117.777797 – ident: e_1_3_6_48_2 doi: 10.1371/journal.pbio.1000365 – volume-title: Geomorph: software for geometric morphometric analyses year: 2021 ident: e_1_3_6_32_2 – volume-title: Morphometrics with R year: 2008 ident: e_1_3_6_35_2 – ident: e_1_3_6_24_2 doi: 10.1007/s00427-020-00645-w – ident: e_1_3_6_57_2 doi: 10.1073/pnas.1409585111 – ident: e_1_3_6_54_2 doi: 10.1016/S0022-5193(84)80050-8 – volume: 53 start-page: 1692 year: 1999 ident: e_1_3_6_11_2 article-title: Mutational contributions to genetic variance–covariance matrices: an experimental approach using induced mutations in Arabidopsis thaliana publication-title: Evolution – ident: e_1_3_6_25_2 doi: 10.1098/rspb.2021.0241 – ident: e_1_3_6_9_2 doi: 10.2307/2410734 – ident: e_1_3_6_21_2 doi: 10.1006/anbe.1999.1342 – ident: e_1_3_6_59_2 doi: 10.1098/rspb.2022.1441 – ident: e_1_3_6_7_2 doi: 10.1534/genetics.118.300995 – ident: e_1_3_6_36_2 doi: 10.1111/jeb.13429 – volume-title: Australia's introduced dung beetles: original release and redistributions year: 1996 ident: e_1_3_6_50_2 – ident: e_1_3_6_27_2 doi: 10.1073/pnas.0809668106 – ident: e_1_3_6_23_2 doi: 10.1111/j.0014-3820.2005.tb01044.x – volume: 29 start-page: 349 year: 1975 ident: e_1_3_6_49_2 article-title: A European dung beetle, Onthophagus taurus Schreber, new to the U.S. (Coleoptera: Scarabaeidae) publication-title: Coleopterists Bullet. doi: 10.5962/p.372750 – ident: e_1_3_6_53_2 doi: 10.1111/j.0014-3820.2005.tb01763.x – ident: e_1_3_6_52_2 doi: 10.1038/nature23473 – ident: e_1_3_6_51_2 doi: 10.1016/j.ympev.2016.08.016 – ident: e_1_3_6_46_2 doi: 10.1016/j.tig.2012.02.002 – ident: e_1_3_6_47_2 doi: 10.1146/annurev.med.60.042307.110802 – start-page: 81 volume-title: Phenotypic plasticity of insects: mechanisms and consequences year: 2009 ident: e_1_3_6_43_2 – ident: e_1_3_6_37_2 doi: 10.1093/sysbio/syt025 – ident: e_1_3_6_18_2 doi: 10.1086/386551 – ident: e_1_3_6_10_2 doi: 10.1098/rspb.2015.1119 – ident: e_1_3_6_30_2 – ident: e_1_3_6_26_2 doi: 10.1002/jez.2413 – ident: e_1_3_6_29_2 doi: 10.1098/rspb.2018.1631 – volume-title: Tpsdig. 2.14 year: 2009 ident: e_1_3_6_31_2 – ident: e_1_3_6_44_2 doi: 10.1111/j.1095-8312.2006.00674.x – ident: e_1_3_6_41_2 doi: 10.1111/evo.14045 – ident: e_1_3_6_2_2 doi: 10.1086/414425 – ident: e_1_3_6_28_2 doi: 10.1073/pnas.1118589109 – volume-title: Clustergeneration: random cluster generation (with specified degree of separation) year: 2020 ident: e_1_3_6_40_2 – ident: e_1_3_6_4_2 doi: 10.1038/ncomms4700 – ident: e_1_3_6_58_2 doi: 10.1098/rspb.2022.1441 |
SSID | ssj0009585 |
Score | 2.4608955 |
Snippet | The degree to which developmental systems bias the phenotypic effects of environmental and genetic variation, and how these biases affect evolution, is subject... |
SourceID | pubmedcentral proquest pubmed crossref |
SourceType | Open Access Repository Aggregation Database Index Database Enrichment Source |
StartPage | 20221441 |
SubjectTerms | Animals Bias Biological Evolution Coleoptera - genetics Evolution Phenotype |
Title | Developmental bias in the evolution and plasticity of beetle horn shape |
URI | https://www.ncbi.nlm.nih.gov/pubmed/36168764 https://www.proquest.com/docview/2718962269 https://pubmed.ncbi.nlm.nih.gov/PMC9515630 |
Volume | 289 |
hasFullText | 1 |
inHoldings | 1 |
isFullTextHit | |
isPrint | |
link | http://utb.summon.serialssolutions.com/2.0.0/link/0/eLvHCXMwnV1ba9swGBVbx2AvY-1u2Q0NBtsIzmxLvj2upV0YpdtDAnkzkiw1ocUudTJYf_0-XSzHWwbdXkxwFNnonHwX6egTQu-YIpBwySogJFIBVTQNCi6igGWScVHFLOJG5XuWTuf06yJZ9CJ2s7tkzSfiZue-kv9BFe4BrnqX7D8g6zuFG_AZ8IUrIAzXW2G8pfiBkeYr1nayRfnDPdgWAoAQWaunrfqCSwk_Hi-b63rcLtnVQAz03Tu0tpMP2BkGJ--cjA8NJVbeaDof2mvm_SYaW_7_otdhTzfG4jf1-TlzHtPofcWNNEZZs652O87cTAQksXpxZtt4gqML9Lqh9S077jmLG9tTgzpqFTnZNqHQs87ydtr3sNB7FiDj5xPdbtI1HBbSPvtWnsxPT8vZ8WJ2F92LIYMIu4kcX4_ZnNbqX8_X88w_DXsfxit_JCG_a2m3gpPZI_TQZRX4s6XIProj6wN034L08wDtOwve4g-uzPjHx-jLgD1YswevagyIY88eDOzBPXtwo7BlD9bswYY9T9D85Hh2NA3csRqBoISug4olecgUZXGooiKRYPQ5rWiaAA6CFIRLQpM8I0pUmY7I45DkIiFZEivFIVsgT9Fe3dTyOcJKUplUOY8Ek5SLHJxlkkrGmLb1acZGKOgGrxSu5rw--uSytNqHvNSDXerBLvVgj9B73_7KVlv5a8u3HRYlGES9ysVq2WzaMoZoq0ghqyhG6JnFxvdF0igF909HKBug5hvoYuvDb-rV0hRdh0xEl9J7cYvnvkQP-r_HK7S3vt7I1xC6rvkbQ8JfaVCceQ |
linkProvider | National Library of Medicine |
openUrl | ctx_ver=Z39.88-2004&ctx_enc=info%3Aofi%2Fenc%3AUTF-8&rfr_id=info%3Asid%2Fsummon.serialssolutions.com&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=Developmental+bias+in+the+evolution+and+plasticity+of+beetle+horn+shape&rft.jtitle=Proceedings+of+the+Royal+Society.+B%2C+Biological+sciences&rft.au=Rohner%2C+Patrick+T&rft.au=Hu%2C+Yonggang&rft.au=Moczek%2C+Armin+P&rft.date=2022-09-28&rft.issn=1471-2954&rft.eissn=1471-2954&rft.volume=289&rft.issue=1983&rft.spage=20221441&rft_id=info:doi/10.1098%2Frspb.2022.1441&rft.externalDBID=NO_FULL_TEXT |
thumbnail_l | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/lc.gif&issn=0962-8452&client=summon |
thumbnail_m | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/mc.gif&issn=0962-8452&client=summon |
thumbnail_s | http://covers-cdn.summon.serialssolutions.com/index.aspx?isbn=/sc.gif&issn=0962-8452&client=summon |