[{"acknowledgement":"This work was supported by the Russian Science Foundation grant N 16-14-10173.","status":"public","quality_controlled":"1","month":"04","volume":524,"publisher":"Elsevier","title":"Two linked loci under mutation-selection balance and Muller’s ratchet","article_processing_charge":"No","department":[{"_id":"GradSch"}],"date_published":"2021-04-24T00:00:00Z","publication_identifier":{"issn":["0022-5193"]},"oa_version":"Preprint","external_id":{"isi":["000659161500002"],"pmid":["33901507"]},"date_updated":"2026-07-06T12:58:31Z","publication_status":"published","_id":"9387","das_tickbox":"1","author":[{"last_name":"Khudiakova","id":"4E6DC800-AE37-11E9-AC72-31CAE5697425","full_name":"Khudiakova, Kseniia","first_name":"Kseniia","orcid":"0000-0002-6246-1465"},{"first_name":"Tatiana Yu.","full_name":"Neretina, Tatiana Yu.","last_name":"Neretina"},{"last_name":"Kondrashov","first_name":"Alexey S.","full_name":"Kondrashov, Alexey S."}],"intvolume":"       524","language":[{"iso":"eng"}],"article_type":"original","publication":"Journal of Theoretical Biology","date_created":"2021-05-12T05:58:42Z","pmid":1,"abstract":[{"text":"We report the complete analysis of a deterministic model of deleterious mutations and negative selection against them at two haploid loci without recombination. As long as mutation is a weaker force than selection, mutant alleles remain rare at the only stable equilibrium, and otherwise, a variety of dynamics are possible. If the mutation-free genotype is absent, generally the only stable equilibrium is the one that corresponds to fixation of the mutant allele at the locus where it is less deleterious. This result suggests that fixation of a deleterious allele that follows a click of the Muller’s ratchet is governed by natural selection, instead of random drift.","lang":"eng"}],"main_file_link":[{"url":"https://www.biorxiv.org/content/10.1101/477489v1","open_access":"1"}],"day":"24","isi":1,"oa":1,"doi":"10.1016/j.jtbi.2021.110729","scopus_import":"1","keyword":["General Biochemistry","Genetics and Molecular Biology","Modelling and Simulation","Statistics and Probability","General Immunology and Microbiology","Applied Mathematics","General Agricultural and Biological Sciences","General Medicine"],"user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","citation":{"short":"K. Khudiakova, T.Y. Neretina, A.S. Kondrashov, Journal of Theoretical Biology 524 (2021).","ama":"Khudiakova K, Neretina TY, Kondrashov AS. Two linked loci under mutation-selection balance and Muller’s ratchet. <i>Journal of Theoretical Biology</i>. 2021;524. doi:<a href=\"https://doi.org/10.1016/j.jtbi.2021.110729\">10.1016/j.jtbi.2021.110729</a>","ista":"Khudiakova K, Neretina TY, Kondrashov AS. 2021. Two linked loci under mutation-selection balance and Muller’s ratchet. Journal of Theoretical Biology. 524, 110729.","apa":"Khudiakova, K., Neretina, T. Y., &#38; Kondrashov, A. S. (2021). Two linked loci under mutation-selection balance and Muller’s ratchet. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/j.jtbi.2021.110729\">https://doi.org/10.1016/j.jtbi.2021.110729</a>","chicago":"Khudiakova, Kseniia, Tatiana Yu. Neretina, and Alexey S. Kondrashov. “Two Linked Loci under Mutation-Selection Balance and Muller’s Ratchet.” <i>Journal of Theoretical Biology</i>. Elsevier, 2021. <a href=\"https://doi.org/10.1016/j.jtbi.2021.110729\">https://doi.org/10.1016/j.jtbi.2021.110729</a>.","ieee":"K. Khudiakova, T. Y. Neretina, and A. S. Kondrashov, “Two linked loci under mutation-selection balance and Muller’s ratchet,” <i>Journal of Theoretical Biology</i>, vol. 524. Elsevier, 2021.","mla":"Khudiakova, Kseniia, et al. “Two Linked Loci under Mutation-Selection Balance and Muller’s Ratchet.” <i>Journal of Theoretical Biology</i>, vol. 524, 110729, Elsevier, 2021, doi:<a href=\"https://doi.org/10.1016/j.jtbi.2021.110729\">10.1016/j.jtbi.2021.110729</a>."},"article_number":"110729","type":"journal_article","year":"2021"},{"user_id":"2DF688A6-F248-11E8-B48F-1D18A9856A87","has_accepted_license":"1","year":"2017","corr_author":"1","type":"journal_article","citation":{"ieee":"T. Priklopil, K. Chatterjee, and M. Nowak, “Optional interactions and suspicious behaviour facilitates trustful cooperation in prisoners dilemma,” <i>Journal of Theoretical Biology</i>, vol. 433. Elsevier, pp. 64–72, 2017.","mla":"Priklopil, Tadeas, et al. “Optional Interactions and Suspicious Behaviour Facilitates Trustful Cooperation in Prisoners Dilemma.” <i>Journal of Theoretical Biology</i>, vol. 433, Elsevier, 2017, pp. 64–72, doi:<a href=\"https://doi.org/10.1016/j.jtbi.2017.08.025\">10.1016/j.jtbi.2017.08.025</a>.","chicago":"Priklopil, Tadeas, Krishnendu Chatterjee, and Martin Nowak. “Optional Interactions and Suspicious Behaviour Facilitates Trustful Cooperation in Prisoners Dilemma.” <i>Journal of Theoretical Biology</i>. Elsevier, 2017. <a href=\"https://doi.org/10.1016/j.jtbi.2017.08.025\">https://doi.org/10.1016/j.jtbi.2017.08.025</a>.","apa":"Priklopil, T., Chatterjee, K., &#38; Nowak, M. (2017). Optional interactions and suspicious behaviour facilitates trustful cooperation in prisoners dilemma. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/j.jtbi.2017.08.025\">https://doi.org/10.1016/j.jtbi.2017.08.025</a>","ista":"Priklopil T, Chatterjee K, Nowak M. 2017. Optional interactions and suspicious behaviour facilitates trustful cooperation in prisoners dilemma. Journal of Theoretical Biology. 433, 64–72.","ama":"Priklopil T, Chatterjee K, Nowak M. Optional interactions and suspicious behaviour facilitates trustful cooperation in prisoners dilemma. <i>Journal of Theoretical Biology</i>. 2017;433:64-72. doi:<a href=\"https://doi.org/10.1016/j.jtbi.2017.08.025\">10.1016/j.jtbi.2017.08.025</a>","short":"T. Priklopil, K. Chatterjee, M. Nowak, Journal of Theoretical Biology 433 (2017) 64–72."},"date_created":"2018-12-11T11:48:16Z","ddc":["000","570"],"publication":"Journal of Theoretical Biology","file":[{"date_updated":"2020-07-14T12:47:58Z","file_name":"2017_JournTheoretBio_Priklopil.pdf","content_type":"application/pdf","file_id":"7047","date_created":"2019-11-19T07:57:39Z","checksum":"4b43af1615ebf1a861840cb03d8a320c","file_size":537323,"creator":"dernst","relation":"main_file","access_level":"open_access"}],"scopus_import":"1","doi":"10.1016/j.jtbi.2017.08.025","oa":1,"page":"64 - 72","isi":1,"day":"21","project":[{"name":"International IST Postdoc Fellowship Programme","grant_number":"291734","_id":"25681D80-B435-11E9-9278-68D0E5697425","call_identifier":"FP7"},{"call_identifier":"FP7","grant_number":"279307","name":"Quantitative Graph Games: Theory and Applications","_id":"2581B60A-B435-11E9-9278-68D0E5697425"}],"abstract":[{"text":"In evolutionary game theory interactions between individuals are often assumed obligatory. However, in many real-life situations, individuals can decide to opt out of an interaction depending on the information they have about the opponent. We consider a simple evolutionary game theoretic model to study such a scenario, where at each encounter between two individuals the type of the opponent (cooperator/defector) is known with some probability, and where each individual either accepts or opts out of the interaction. If the type of the opponent is unknown, a trustful individual accepts the interaction, whereas a suspicious individual opts out of the interaction. If either of the two individuals opt out both individuals remain without an interaction. We show that in the prisoners dilemma optional interactions along with suspicious behaviour facilitates the emergence of trustful cooperation.","lang":"eng"}],"pmid":1,"file_date_updated":"2020-07-14T12:47:58Z","intvolume":"       433","author":[{"first_name":"Tadeas","full_name":"Priklopil, Tadeas","last_name":"Priklopil","id":"3C869AA0-F248-11E8-B48F-1D18A9856A87"},{"orcid":"0000-0002-4561-241X","first_name":"Krishnendu","full_name":"Chatterjee, Krishnendu","id":"2E5DCA20-F248-11E8-B48F-1D18A9856A87","last_name":"Chatterjee"},{"last_name":"Nowak","first_name":"Martin","full_name":"Nowak, Martin"}],"das_tickbox":"1","license":"https://creativecommons.org/licenses/by-nc-nd/4.0/","_id":"744","article_type":"original","tmp":{"name":"Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International (CC BY-NC-ND 4.0)","image":"/images/cc_by_nc_nd.png","legal_code_url":"https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode","short":"CC BY-NC-ND (4.0)"},"language":[{"iso":"eng"}],"volume":433,"month":"11","quality_controlled":"1","status":"public","publication_status":"published","date_updated":"2026-07-07T13:11:54Z","external_id":{"isi":["000412039800007"],"pmid":["28867224"]},"oa_version":"Submitted Version","ec_funded":1,"department":[{"_id":"KrCh"}],"publication_identifier":{"issn":["0022-5193"]},"date_published":"2017-11-21T00:00:00Z","article_processing_charge":"No","publist_id":"6923","title":"Optional interactions and suspicious behaviour facilitates trustful cooperation in prisoners dilemma","publisher":"Elsevier"},{"external_id":{"pmid":["16098538 "]},"date_updated":"2026-08-27T09:50:57Z","publication_status":"published","publication_identifier":{"issn":["0022-5193"]},"date_published":"2006-01-01T00:00:00Z","oa_version":"None","publist_id":"2461","article_processing_charge":"No","publisher":"Elsevier","title":"Determinants of simulated RNA evolution.","volume":238,"month":"01","status":"public","article_type":"original","OA_type":"closed access","language":[{"iso":"eng"}],"intvolume":"       238","author":[{"id":"2BB22BC2-F248-11E8-B48F-1D18A9856A87","last_name":"Kupczok","first_name":"Anne","full_name":"Kupczok, Anne"},{"first_name":"Peter","full_name":"Dittrich, Peter","last_name":"Dittrich"}],"_id":"3767","doi":"10.1016/j.jtbi.2005.06.019","page":"726 - 35","day":"01","abstract":[{"text":"Models of RNA secondary structure folding are widely used to study evolution in theory and simulation. However, systematic studies of the parameters involved are rare. In this paper, we study by simulation how RNA evolution is influenced by three different factors, namely the mutation rate, scaling of the fitness function, and distance measure. We found that for low mutation rates the qualitative evolutionary behavior is robust with respect to the scaling of the fitness function. For efficient mutation rates, which are close to the error threshold, scaling and distance measure have a strong influence on the evolutionary behavior. A global distance measure that takes sequence information additively into account lowers the error threshold. When using a local sequence-structure alignment for the distance, we observed a smoother evolution of the fitness over time. Finally, in addition to the well known error threshold, we identify another threshold of the mutation rate, called divergence threshold, where the qualitative transient behavior changes from a localized to an exploratory search.","lang":"eng"}],"pmid":1,"issue":"3","date_created":"2018-12-11T12:05:03Z","publication":"Journal of Theoretical Biology","extern":"1","type":"journal_article","year":"2006","citation":{"ieee":"A. Kupczok and P. Dittrich, “Determinants of simulated RNA evolution.,” <i>Journal of Theoretical Biology</i>, vol. 238, no. 3. Elsevier, pp. 726–35, 2006.","mla":"Kupczok, Anne, and Peter Dittrich. “Determinants of Simulated RNA Evolution.” <i>Journal of Theoretical Biology</i>, vol. 238, no. 3, Elsevier, 2006, pp. 726–35, doi:<a href=\"https://doi.org/10.1016/j.jtbi.2005.06.019\">10.1016/j.jtbi.2005.06.019</a>.","chicago":"Kupczok, Anne, and Peter Dittrich. “Determinants of Simulated RNA Evolution.” <i>Journal of Theoretical Biology</i>. Elsevier, 2006. <a href=\"https://doi.org/10.1016/j.jtbi.2005.06.019\">https://doi.org/10.1016/j.jtbi.2005.06.019</a>.","apa":"Kupczok, A., &#38; Dittrich, P. (2006). Determinants of simulated RNA evolution. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/j.jtbi.2005.06.019\">https://doi.org/10.1016/j.jtbi.2005.06.019</a>","ista":"Kupczok A, Dittrich P. 2006. Determinants of simulated RNA evolution. Journal of Theoretical Biology. 238(3), 726–35.","ama":"Kupczok A, Dittrich P. Determinants of simulated RNA evolution. <i>Journal of Theoretical Biology</i>. 2006;238(3):726-735. doi:<a href=\"https://doi.org/10.1016/j.jtbi.2005.06.019\">10.1016/j.jtbi.2005.06.019</a>","short":"A. Kupczok, P. Dittrich, Journal of Theoretical Biology 238 (2006) 726–35."},"user_id":"317138e5-6ab7-11ef-aa6d-ffef3953e345"},{"_id":"3656","author":[{"last_name":"Rouhani","first_name":"Shahin","full_name":"Rouhani, Shahin"},{"id":"4880FE40-F248-11E8-B48F-1D18A9856A87","last_name":"Barton","orcid":"0000-0002-8548-5240","full_name":"Barton, Nicholas H","first_name":"Nicholas H"}],"intvolume":"       126","language":[{"iso":"eng"}],"article_type":"original","status":"public","quality_controlled":"1","month":"05","volume":126,"title":"The probability of peak shifts in a founder population","publisher":"Elsevier","article_processing_charge":"No","publist_id":"2727","publication_identifier":{"eissn":["1095-8541"],"issn":["0022-5193"]},"date_published":"1987-05-07T00:00:00Z","oa_version":"None","date_updated":"2022-02-04T12:36:32Z","publication_status":"published","user_id":"ea97e931-d5af-11eb-85d4-e6957dddbf17","citation":{"ieee":"S. Rouhani and N. H. Barton, “The probability of peak shifts in a founder population,” <i>Journal of Theoretical Biology</i>, vol. 126, no. 1. Elsevier, pp. 51–62, 1987.","mla":"Rouhani, Shahin, and Nicholas H. Barton. “The Probability of Peak Shifts in a Founder Population.” <i>Journal of Theoretical Biology</i>, vol. 126, no. 1, Elsevier, 1987, pp. 51–62, doi:<a href=\"https://doi.org/10.1016/S0022-5193(87)80100-5\">10.1016/S0022-5193(87)80100-5</a>.","chicago":"Rouhani, Shahin, and Nicholas H Barton. “The Probability of Peak Shifts in a Founder Population.” <i>Journal of Theoretical Biology</i>. Elsevier, 1987. <a href=\"https://doi.org/10.1016/S0022-5193(87)80100-5\">https://doi.org/10.1016/S0022-5193(87)80100-5</a>.","apa":"Rouhani, S., &#38; Barton, N. H. (1987). The probability of peak shifts in a founder population. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/S0022-5193(87)80100-5\">https://doi.org/10.1016/S0022-5193(87)80100-5</a>","ista":"Rouhani S, Barton NH. 1987. The probability of peak shifts in a founder population. Journal of Theoretical Biology. 126(1), 51–62.","ama":"Rouhani S, Barton NH. The probability of peak shifts in a founder population. <i>Journal of Theoretical Biology</i>. 1987;126(1):51-62. doi:<a href=\"https://doi.org/10.1016/S0022-5193(87)80100-5\">10.1016/S0022-5193(87)80100-5</a>","short":"S. Rouhani, N.H. Barton, Journal of Theoretical Biology 126 (1987) 51–62."},"type":"journal_article","year":"1987","extern":"1","publication":"Journal of Theoretical Biology","issue":"1","date_created":"2018-12-11T12:04:28Z","abstract":[{"text":"We have analysed the role of sampling drift in inducing shifts between alternative adaptive peaks, in small and rapidly growing populations. Using a simple model of disruptive selection on a polygenic character, we calculate the net probabilityofapeakshift. If the growth rate is high, theprobabilityofashiftina growing population is insensitive to selection on the character. Assuming that the character is effectively neutral during the brief initial increase, we find that theprobabilityofapeakshift is given by theprobabilityof finding a standard normal variate greater than √2ΔV where ΔV is the reduction in additive genetic variance during the growth period. This result holds for arbitrary pattern of increase in size, provided that the rate of increase is high enough for selection to be negligible, and the character depends on a large number of loci. Comparing theprobabilityofpeakshiftsin founding populations with the rate ofshiftsin static and allopatric populations it appears that although strongly selected shifts are only likely to occur ina growing population, a static population is a more congenial setting for adaptive shifts.","lang":"eng"}],"main_file_link":[{"url":"https://www.sciencedirect.com/science/article/pii/S0022519387801005?via%3Dihub"}],"day":"07","page":"51 - 62","scopus_import":"1","doi":"10.1016/S0022-5193(87)80100-5"},{"citation":{"ama":"Barton NH, Rouhani S. The frequency of shifts between alternative equilibria. <i>Journal of Theoretical Biology</i>. 1987;125(4):397-418. doi:<a href=\"https://doi.org/10.1016/S0022-5193(87)80210-2\">10.1016/S0022-5193(87)80210-2</a>","short":"N.H. Barton, S. Rouhani, Journal of Theoretical Biology 125 (1987) 397–418.","ista":"Barton NH, Rouhani S. 1987. The frequency of shifts between alternative equilibria. Journal of Theoretical Biology. 125(4), 397–418.","mla":"Barton, Nicholas H., and Shahin Rouhani. “The Frequency of Shifts between Alternative Equilibria.” <i>Journal of Theoretical Biology</i>, vol. 125, no. 4, Elsevier, 1987, pp. 397–418, doi:<a href=\"https://doi.org/10.1016/S0022-5193(87)80210-2\">10.1016/S0022-5193(87)80210-2</a>.","ieee":"N. H. Barton and S. Rouhani, “The frequency of shifts between alternative equilibria,” <i>Journal of Theoretical Biology</i>, vol. 125, no. 4. Elsevier, pp. 397–418, 1987.","apa":"Barton, N. H., &#38; Rouhani, S. (1987). The frequency of shifts between alternative equilibria. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/S0022-5193(87)80210-2\">https://doi.org/10.1016/S0022-5193(87)80210-2</a>","chicago":"Barton, Nicholas H, and Shahin Rouhani. “The Frequency of Shifts between Alternative Equilibria.” <i>Journal of Theoretical Biology</i>. Elsevier, 1987. <a href=\"https://doi.org/10.1016/S0022-5193(87)80210-2\">https://doi.org/10.1016/S0022-5193(87)80210-2</a>."},"language":[{"iso":"eng"}],"type":"journal_article","year":"1987","article_type":"original","_id":"3661","user_id":"ea97e931-d5af-11eb-85d4-e6957dddbf17","author":[{"full_name":"Barton, Nicholas H","first_name":"Nicholas H","orcid":"0000-0002-8548-5240","last_name":"Barton","id":"4880FE40-F248-11E8-B48F-1D18A9856A87"},{"last_name":"Rouhani","first_name":"Shahin","full_name":"Rouhani, Shahin"}],"intvolume":"       125","title":"The frequency of shifts between alternative equilibria","publisher":"Elsevier","abstract":[{"lang":"eng","text":"We derive a formula giving thefrequency with which random drift shifts a population betweenalternativeequilibria. This formula is valid when such shifts are rare (Ns &gt;&gt; 1), and applies over a wide range of mutation rates. When the number of mutations entering the population is low (4Nμ &lt;&lt; 1), the rate of stochastic shifts reduces to the product ofthe mutation rate and the probability of fixation of a single mutation. However, when many mutations enter the population in each generation (4Nμ &gt;&gt; 1), the rate is higher than would be expected if mutations were established independently, and converges to that given by a gaussian approximation. We apply recent results on bistable systems to extend this formula to the general multidimensional case. This gives an explicit expression for thefrequencyof stochastic shifts, which depends only on theequilibrium probability distribution near the saddle point separating thealternative stable states. The plausibility of theories of speciation through random drift are discussed in the light of these results."}],"publist_id":"2722","article_processing_charge":"No","main_file_link":[{"url":"https://www.sciencedirect.com/science/article/pii/S0022519387802102?via%3Dihub"}],"publication_identifier":{"eissn":["1095-8541"],"issn":["0022-5193"]},"page":"397 - 418","date_published":"1987-04-21T00:00:00Z","day":"21","oa_version":"None","date_updated":"2022-02-04T10:39:15Z","publication_status":"published","doi":"10.1016/S0022-5193(87)80210-2","extern":"1","publication":"Journal of Theoretical Biology","status":"public","month":"04","issue":"4","volume":125,"date_created":"2018-12-11T12:04:29Z"},{"citation":{"apa":"Barton, N. H., &#38; Post, R. J. (1986). Sibling competition and the advantage of mixed families. <i>Journal of Theoretical Biology</i>. Elsevier. <a href=\"https://doi.org/10.1016/S0022-5193(86)80033-9\">https://doi.org/10.1016/S0022-5193(86)80033-9</a>","chicago":"Barton, Nicholas H, and R.J. Post. “Sibling Competition and the Advantage of Mixed Families.” <i>Journal of Theoretical Biology</i>. Elsevier, 1986. <a href=\"https://doi.org/10.1016/S0022-5193(86)80033-9\">https://doi.org/10.1016/S0022-5193(86)80033-9</a>.","ieee":"N. H. Barton and R. J. Post, “Sibling competition and the advantage of mixed families,” <i>Journal of Theoretical Biology</i>, vol. 120, no. 4. Elsevier, pp. 381–387, 1986.","mla":"Barton, Nicholas H., and R. J. Post. “Sibling Competition and the Advantage of Mixed Families.” <i>Journal of Theoretical Biology</i>, vol. 120, no. 4, Elsevier, 1986, pp. 381–87, doi:<a href=\"https://doi.org/10.1016/S0022-5193(86)80033-9\">10.1016/S0022-5193(86)80033-9</a>.","ama":"Barton NH, Post RJ. Sibling competition and the advantage of mixed families. <i>Journal of Theoretical Biology</i>. 1986;120(4):381-387. doi:<a href=\"https://doi.org/10.1016/S0022-5193(86)80033-9\">10.1016/S0022-5193(86)80033-9</a>","short":"N.H. Barton, R.J. Post, Journal of Theoretical Biology 120 (1986) 381–387.","ista":"Barton NH, Post RJ. 1986. Sibling competition and the advantage of mixed families. Journal of Theoretical Biology. 120(4), 381–387."},"year":"1986","type":"journal_article","user_id":"ea97e931-d5af-11eb-85d4-e6957dddbf17","abstract":[{"lang":"eng","text":"It is noted that the sibling competition model for the evolution of sex and recombination, as it has been developed so far, involves truncation selection. After briefly reviewing aspects of the development and behaviour of such models an analytical treatment is presented which involves additive selection. Additive selection, as compared with truncation selection, decreases the advantage of sex to such an extent that it is unlikely that sibling competition could overcome its intrinsic two-fold cost, although it could still be important in promoting family variability produced by other mechanisms, such as polyandry."}],"page":"381 - 387","day":"21","doi":"10.1016/S0022-5193(86)80033-9","extern":"1","publication":"Journal of Theoretical Biology","issue":"4","date_created":"2018-12-11T12:08:15Z","language":[{"iso":"eng"}],"article_type":"original","_id":"4323","author":[{"id":"4880FE40-F248-11E8-B48F-1D18A9856A87","last_name":"Barton","orcid":"0000-0002-8548-5240","full_name":"Barton, Nicholas H","first_name":"Nicholas H"},{"first_name":"R.J.","full_name":"Post, R.J.","last_name":"Post"}],"intvolume":"       120","title":"Sibling competition and the advantage of mixed families","publisher":"Elsevier","article_processing_charge":"No","publist_id":"1720","oa_version":"None","date_published":"1986-06-21T00:00:00Z","publication_identifier":{"issn":["0022-5193"],"eissn":["1095-8541"]},"publication_status":"published","date_updated":"2022-01-31T14:44:50Z","acknowledgement":"We would like to thank M. Bulmer for his helpful comments. R. J. Post was supported during this work by an MRC Postdoctoral Fellowship, and N. Barton by an SRC Postdoctoral Fellowship. \r\n","quality_controlled":"1","status":"public","month":"06","volume":120}]