Official withdraw from the course is only possible if the student attends neither the final theoretical nor the practical exam.

The final grade of the students who did not reach the minimum grade to pass the course in the practical or the objective test, but whose cumulative score happened to be higher than 50, will be a 4.9 (FAILED).

In the second opportunity, the same evaluation methodology will be used as in the first one.

In the event that a student, for duly justified reasons, cannot attend the official exams of the subject, he/she will be examined orally. If he/she is unable to take the continuous evaluation tests, or if he/she does not obtain the maximum possible points with these tests, he/she may take an additional block of exercises in the official exam, in order to recover the points lost.

For the computation of the final grade of students with recognition of part-time dedication and academic dispensation of attendance, both in the opportunity of the end of term and in the second opportunity, the grade obtained in the theoretical exam and the corresponding practical part (see above format of both exams) will be taken into account, representing 75% and 25% of the final grade, respectively.

The fraudulent performance of the evaluation tests or activities will directly imply the grade of FAILED (0) in the subject at the corresponding opportunity.

Learning outcomes	Study programme competences
Capacity to interpret and to analyze the biological problems, as well as the human nature itself, from an evolutionary perspective	A7 A12 A18 A21	B1 B2 B3 B4 B5 B6 B7
Choice of the techniques and methods more adequate to tackle the study of a specific evolutionary problem	A7 A12 A18 A24	B1 B2 B3 B4 B5 B6 B7
Use of the genetic information to manage, to preserve and to restore populations.	A7 A12 A18 A21 A24 A27	B1 B2 B3 B4 B5 B6 B7

Topic	Sub-topic
OVERVIEW OF EVOLUTIONARY BIOLOGY	Brief history of Evolutionary Biology. Population genetics. Molecular evolutionary genetics. Evolutionary biology of development (evo-devo). Evolutionary genomics. The National Center for Biotechnology Information (NCBI) databases. Genome browsers (NCBI, UCSC, Ensembl). International projects to sample human genomes (IGSR, varsome)
MACROEVOLUTION	Evolution above the species level. Timeline of life on earth. The three domains of life. Using phylogenies to reconstruct the deep past. Diversification of eukaryotes. The species concept in paleontology. Patterns of macroevolution. Mass extinctions. Differences among clades in species diversity. The evolution of complex biological structures through the fossil record.
THE BUILDING OF EVOLUTIONARY MODULES	Promiscuous proteins; molecular machines; modular evolution of proteins. Evolutionary tinkering. Biochemical construction kits. Adaptations, exaptations and spandrels. Evolution of developmental programs: recycling networks. Retrograde and intercalary evolution. Gene duplications. Recruitment. Horizontal transmission. Linkage groups. Randomization effect of recombination. Genetic coadaptation. Supergenes.
MOLECULAR PHYLOGENIES	Cladograms and phylograms. Coalescence theory. Monophyletic, paraphyletic and polyphyletic taxa. Gene trees and species trees. Methods of molecular phylogenetics. The human evolutionary tree.
THE ORIGINS OF SPECIES	Concepts of species. Main questions related to speciation. Intrinsic reproductive barriers of isolation. Speciation and fitness landscapes: the shifting-balance theory. Modes of speciation. Adaptive radiations. Magic traits. Evolution of hybrid genetic incompatibilities. General rules of speciation and evolutionary diversification. Phyletic and cladistic evolution in the fossil record.
QUANTITATIVE GENETICS	Continuous, discontinuous and threshold characters. Breeding value and genotypic value of a genotype. Environmental value. Environmental sensitivity of a genotype. Components of phenotypic variance. Heritability. Estimation of the minimum number of loci underlying a quantitative trait (QTL). Mapping of QTLs. Genome-wide association studies (GWAS).
CONSEQUENCES OF REPRODUCTIVE SYSTEMS AND TYPES OF MATING ON THE ORGANIZATION OF GENETIC VARIATION	Maintenance of genetic variation in populations with sexual reproduction and random mating: Hardy-Weinberg law (H-W); deviations from H-W expectations. Effects of asexual reproduction and non-random mating on genotype frequencies: parthenogenesis; self-fertilization; inbreeding and relatedness coefficients; regular systems of inbreeding; phenotypic assortative mating. Genetic admixture.
RANDOM GENETIC CHANGES IN POPULATIONS OF SMALL SIZE	Sampling of gametes and random walk of gene frequencies. Wright-Fisher model. Dispersion of gene frequencies among subpopulations. Rate of fixation within subpopulations and genomes. Effective population size. Founder effects and population bottlenecks. Wahlund effect.
MUTATION AND MIGRATION	Classes of mutations: nucleotide substitutions; insertions and deletions; duplications; chromosome rearrangements. Mutation rates. Change in gene frequency due to mutation. The fate of a single mutant. Models of mutation in molecular population genetics. Migration and gene flow. Change in gene frequency due to migration; the island model. Mutation and migration in finite populations.
EFFECTS OF NATURAL SELECTION ON PHENOTYPES AND GENE FREQUENCIES	Natural selection. Biological fitness. Types of selection. Selection on quantitative traits. Measuring multivariate selection. Selection on correlated characters. Case study: the genetic basis of adaptation to high altitude in humans. Good genes or bad genes? Haploid and diploid basic models of selection. Polymorphisms maintained by constant selection coefficients. Fitness estimation. Fitness landscapes.
POLYMORPHISMS MAINTAINED BY VARYING SELECTION COEFFICIENTS	Spatial and temporal fitness variation: coarse-grained and fine-grained environments. Endocyclic selection. Trade-offs between fitness components. Antagonistic pleiotropy. Frequency-dependent selection. Cooperation, altruism and kin-selection.
COMBINED ACTION OF SELECTION AND OTHER EVOLUTIONARY FORCES. VARYING SELECTION COEFFICIENTS	Mutation-selection balance. Genetic load of populations. The role of recombination: Muller's ratchet and the degeneration of Y chromosomes; Hill-Robertson effects. Evolution of sex chromosomes. Equilibrium between selection and gene flow; gene clines. Selection in finite populations: neutral, nearly-neutral and selected mutations.
ENGINES OF EVOLUTION	Red Queen dynamics. Interspecies antagonisms. Sexual conflicts. Sexual selection vs. natural selection. Parent-offspring conflicts. Intergenomic conflicts: cytoplasmic incompatibility. Intragenomic conflicts: selfish genetic elements.
THE EVOLUTION OF SEX DETERMINATION	What is meiotic sex? The costs and benefits of sex. The diversity of sexual cycles among eukaryotes. Molecular mechanisms of sex determination. Sex determination in angiosperms. Sex determination in animals. Self-incompatibility systems. Quantitative genetics of sex determination: genotypic versus environmental sex determination. Systems lacking differentiated sex chromosomes. Transitions among sex-determination systems.
THE NEUTRAL THEORY OF MOLECULAR EVOLUTION. MOLECULAR FOOTPRINTS OF NATURAL SELECTION	The neutral theory of molecular evolution. Molecular clocks. Models of DNA evolution. Limits of nucleotide divergence. Estimates of the number of nucleotide substitutions. Substitution rates. Pseudogenes. Direct effects of selection on nucleotide polymorphism and divergence. The importance of recombination: selective sweep and background selection. Selection and demographic history can leave similar footprints on DNA variation. Statistical tests.

Methodologies / tests	Competencies	Ordinary class hours	Student’s personal work hours	Total hours
Introductory activities	B1 B4 B5 B6	1	0.5	1.5
Guest lecture / keynote speech	A7 A12 A18 A24 B1 B3 B4 B6	21	63	84
Problem solving	A7 A27 B1 B2 B3 B4 B5 B7	7	24.5	31.5
ICT practicals	A7 A21 B2 B4	14	14	28
Objective test	A7 A12 A18 A21 A24 B1 B2	4	0	4

Personalized attention		1	0	1

(*)The information in the planning table is for guidance only and does not take into account the heterogeneity of the students.

Methodologies	Description
Introductory activities	Teacher: Presents the teaching guide of the subject, clarifies doubts, organizes the students for the activities. Student: Takes notes, formulates doubts and questions.
Guest lecture / keynote speech	Teacher: Explains the theoretical foundations. Student: Observes, assimilates and takes notes. Formulate doubts and questions. Memorizes. Reads the recommended texts.
Problem solving	Teacher: Formulate problems and provide guidance for their resolution. Student: Works individually or in groups, looks for information and solves the formulated questions.
ICT practicals	Teacher. - Presents the objectives, prepares the material and equipment, explains the methods, provides a script, assists the students. Student. - Experiments, analyzes and prepares a report.
Objective test	Teacher. - Asks questions and evaluates student responses. Student. - He consults his support materials and answers the questions.

Methodologies	Competencies	Description	Qualification
Problem solving	A7 A27 B1 B2 B3 B4 B5 B7	Resolution in the classroom of calculation exercises complementary to the theoretical classes.	15
ICT practicals	A7 A21 B2 B4	Practical exercises of bioinformatics. Compulsory: to avoid failing the subject, every student should obtain at least 15 points in this exam.	25
Objective test	A7 A12 A18 A21 A24 B1 B2	Set of exercises and questions of different types (multiple choice, short answer, complete, association, etc.) related to any of the contents of the syllabus. The test is developed in two phases. The first one is not face-to-face, and consists of a series of questionnaires on the Moodle platform, which must be answered on dates and times set in advance throughout the course. The contribution of this phase to the test is a maximum of 25 points. The second phase, which corresponds to the official exam of the subject, is face-to-face and consists of a series of calculation exercises and multiple choice test questions. The cumulative contribution of the two phases to the final grade of the subject is a maximum of 60 points. Compulsory: to avoid failing the subject, every student should obtain at least 35 points in this test to pass the subject.	60

Assessment comments
Official withdraw from the course is only possible if the student attends neither the final theoretical nor the practical exam. The final grade of the students who did not reach the minimum grade to pass the course in the practical or the objective test, but whose cumulative score happened to be higher than 50, will be a 4.9 (FAILED). In the second opportunity, the same evaluation methodology will be used as in the first one. In the event that a student, for duly justified reasons, cannot attend the official exams of the subject, he/she will be examined orally. If he/she is unable to take the continuous evaluation tests, or if he/she does not obtain the maximum possible points with these tests, he/she may take an additional block of exercises in the official exam, in order to recover the points lost. For the computation of the final grade of students with recognition of part-time dedication and academic dispensation of attendance, both in the opportunity of the end of term and in the second opportunity, the grade obtained in the theoretical exam and the corresponding practical part (see above format of both exams) will be taken into account, representing 75% and 25% of the final grade, respectively. The fraudulent performance of the evaluation tests or activities will directly imply the grade of FAILED (0) in the subject at the corresponding opportunity.

Basic	Cutter, A. D. (2019). A primer of molecular population genetics. OUP Oxford Hartl, D. L. (2020). A primer of population genetics and genomics. OUP Oxford Futuyma, D. J., and Kirkpatrick, M. (2017). Evolution. Sinauer Associates Zimmer, C. and Emlen, D. (2015). Evolution: Making sense of life. Roberts and Company Publishers Herron, J. D., and Freeman, S. (2014). Evolutionary Analysis. . Pearson Caballero, A. (2017). Genética Cuantitativa. Síntesis Hedrick, P.W. (2011). Genetics of Populations.. Jones & Bartlett Hahn, M. W. (2018). Molecular Population Genetics. OUP USA DeSalle, R. (2013). Phylogenomics: A primer. Routledge Lane, N (2018). Power, Sex, Suicide. OUP Oxford Beukeboom, L., and Perrin, N. (2014). The evolution of sex determination. OUP Oxford Shubin, N. (2015). Tu pez interior. Capitán Swing

Complementary	Sampedro, J. (2007). Deconstruyendo a Darwin: Los Enigmas de la Evolución a la Luz de la Nueva Genética.. Síntesis Fontdevila, A., y Moya, A. (2003). Evolución. Origen, adaptación y divergencia de las especies.. Síntesis Barton, N. (2007). Evolution. Cold Spring Harbor Lab. Press. Ridley, M. (2004). Evolution. Blackwell Avise, J. C. (2006). Evolutionary Pathways in Nature. A Phylogenetic Approach. . Cambridge Univ. Press. Fontdevila, A., y Moya, A. (1999). Introducción a la genética de poblaciones. Síntesis Bromham, L. (2008). Reading the Story in DNA: A Beginners Guide to Molecular Evolution. . Oxford Univ. Press. Coyne, J. A. (2009). Why Evolution is True. Viking