Blaustein Institutes for Desert Research

Albert Katz International School for Desert Studies


Institute for Dryland Environmental Research

Mitrani Department of Desert Ecology




Paul Goldstein and Lillian Goldstein Lande Scientific Exchange Fund


Ben-Gurion University of the Negev







Camp Evolution IV


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Last updated April 8, 2008



Sede-Boqer Campus, April 13-17, 2008


Organizer: Ariel Novoplansky



BEWARE! The number of participants is limited. Registration deadline: March 15, 2008





The workshop will feature a lecture series by Alexey Kondrashov, Life Sciences Institute, University of Michigan, Ann Arbor on

Unsolved Problems in Evolutionary Biology


as well as presentations by other participants.


Tentative syllabus


        Prologue I how do we know that modern life is a product of past evolution?

Indirect and direct evidence of past evolution. Suboptimality, homology, unforced hierarchy, patterns in spatial distribution of life, scenario- and theory-based evidence. What can we learn about evolution from past history of life?


        Prologue II are there any unsolved problems in phylogenetic reconstructions?

Reconstructing constant rate evolution. Reconstructing exclusively divergent evolution. Detecting variations in rates of evolution. Detecting homoplasy. Are there any inherently homoplasy-free traits? What phylogenetic problems will remain unresolved five years from now?


        Unsolved problem I: the roles of negative and positive selection

Review of theory of natural selection. Classification of modes of selection. Action of selection in the simplest case Fisher's Fundamental Theorem and dynamics of allele replacement. Methods of detecting negative, positive, and balancing selection. Data on negative and positive selection. Within-population variation and macroevolution.


        Unsolved problem II: the nature of fitness landscapes

Spaces of genotypes and of phenotypes. Simple models of fitness landscapes. Types of epistasis: narrowing, widening, sign, multidimensional. Epistasis at different scales. Data on fitness landscapes. Genotype-phenotype maps. What are real phenotypic targets of natural selection?


        Unsolved problem III: is evolution mutation-limited or selection-limited?

Evidence for mutation-limited evolution: frequencies of replacements of different kinds, gene duplications, domestication of transposable elements. Evidence for selection-driven evolution: stasis, episodes of rapid evolution. Can the answer be different at different levels of organisation? Gillespie-Orr's model of adaptation. Any positive role for counteradaptive changes? Predictability of evolution.


        Unsolved problem IV: evolution of sex.

Review of sex in nature. Immediate benefit vs. variation and selection. 2x2 classification of possible advantages of sex. Long-term and short-term effects. Degeneration of non-recombining chromosomes. Environmental explanations and their difficulties. Mutational explanations and their difficulties. Data on mutation rates and epistasis. Problems with facultative sex and crossing-over.


        Unsolved problem V: how can new species evolve?

Similarity, relatedness, connectedness, compatibility. Dobzhansky-Muller incompatibility. Phyletic speciation. Orr's snowball. Haldane's rule. Data on incompatibilities between species. Coevolution and the role of positive selection. Data on sympatric speciation. Disruptive selection and incompatibility selection.


        Unsolved problem VI: How can altruism evolve?

Individual selection and group selection. Kin selection. Inclusive fitness. Evolutinarily stable strategies. Direct, indirect and network reciprocity. Cooperation, Prisoner's dilemma. Altruistic punishment. Eusociality. Multicellularity and coloniality.


        Unsolved problem VII: How can we study the evolution of complex phenotypes?

Generalizations vs. theories. Complexity, designability, modularity, robustness. Are there any general laws that govern fitness landscapes of complex phenotypes? Models and experiments on the evolution of molecules. Models of the evolution of cells and of multicellular organisms.

        Unsolved problem VIII: How did life appear?

What is life? What do we know time interval, conditions, prebiotic organic synthesis, LUCA. How probable is the origin of life? Replicators and reproducers. Hypercycle. RNA world, primitive cells, or both? Experimental data.


The workshop's language will be English.


Getting to Midreshet Ben-Gurion and the Albert Katz International School for Desert Studies






Sun, April 13

Mon, April 14

Tues, April 15

Wed, April 16

Thurs, April 17

08:00 08:20


Arrival and






08:30 10:15

Alex Kondrashov

Negative and positive selection

Alex Kondrashov

Mutation and selection limits to evolution

Alex Kondrashov


Alex Kondrashov

Complex phenotypes

10:15 10:45

Coffee break

Coffee break

Coffee break

Coffee break

10:45 12:30

Alex Kondashov

Adaptive landscapes

Alex Kondashov

Evolution of sex

Alex Kondashov


Alex Kondashov

Emergence of life

12:30 13:30





14:00 18:00

Alex Kondrashov

Prologue I













Coffee break

Alex Kondashov: Prologue II


19:00 20:00








Dinner Party


20:30 21:30

Michal Segoli


The evolution of polyembryony in parasitoid wasps

Nadav Kashtan


Evolution in varying environments: rapid emergence of modular systems

Ofir Cohen


A Bayesian Framework to Analyze Phyletic Patterns

Eyal Privman


Selection constraints on selfish homing endonucleases facilitate their use in gene therapy


21:30 - ??