Summer 2016 Positions Available: Benson-Amram Zebra Finch Lab Behavioral Genetics 2 Goal: To investigate personality and problem-solving ability in individual and pairs of captive zebra finches in Dr. Sarah Benson-Amram’s Animal Behavior & Cognition Lab.

Responsibilities: Assist with planning and carrying out behavioral experiments with zebra finches. Help maintain routine husbandry. Extract data from video-recorded experimental trials. Qualities Required of Applicants: Interest in animal behavior and cognition Dedication to the project. Ability to work odd hours and be flexible with scheduling (this is not a typical 9-5 job!). Ability to work well with others and independently. Ability to maintain a positive attitude. Organization skills and proficiency in use of Microsoft Office.

Benefits: Students can earn 1-2 credits for completion of summer work. Successful volunteers will be encouraged to apply for an EPSCoR grant in the fall to continue working with the Benson-Amram lab. Successful volunteers can use this experience on their CV/resume and request letters of recommendation in the future. Hands-on research experience.

To be considered for this position, interested candidates must submit the following to Lisa Barrett ([email protected]) no later than* March 11th:

1) Cover letter or letter of interest 2) CV or Resume 3) 3 references (name and contact information)

*If you are ready to begin this position before Summer 2016, please let us know and we will get you started!

Experimental manipulation of gene function: knockout studies • Knockout technique – Procedure that eliminates the expression of a gene

• single gene-effects fosB +

• insert a non-functional gene sequence, compare the resulting phenotype fosB - to the ‘wild-type’ Social environment and gene expression in birds

• Research question: What is the role of the FoxP2 gene in song development? (Haesler et al. 2007) • Song system – Area of avian brain that controls song production

Social environment and gene expression in birds Social environment and gene expression in birds

• Methods: – Zebra finches (Taeniopygia guttata) – Knockdown technique – used a virus to Spectrogram: insert short sections of RNA into FoxP2 gene to reduce its expression Allows researchers – Controls had short sections of RNA to characterize placed in a noncoding region of DNA acoustic structure – Adult male tutors and young juvenile of vocalizations males housed together – Recorded songs Social environment and gene expression in birds

• Results: – Knockdown birds had much lower FoxP2 expression than controls

Social environment and gene expression in birds Microarray Analysis

• Results: – Knockdown birds tended to omit specific syllables in their songs • Conclusion: – FoxP2 is required for normal song development QTL mapping to identify genes associated QTL mapping with behavior

• Quantitative trait loci (QTL) • Use marker loci that are easily assayed, but causally – Stretches of DNA that either contain or are linked to genes unrelated to the trait in question to identify the influencing a trait such as behavior approximate locations of the unknown alleles that • QTL mapping affect the behavioral trait of interest – Statistical technique that combines genetic information with trait information to determine which regions of the genome contain the genes that influence the trait QTLs • Candidate genes – Major genes suspected of contributing to a large amount of the phenotypic variation in a specific trait QTL mapping • Step 1: Select two parental strains that a) differ considerably in their values of the trait of interest and b) differ at a set of marker alleles

Genes and Environment

• Have focused on genetic influences on behavior. • However, variation in behavior among individuals results from both variation in gene alleles and variation in environments. • How can we characterize the effects of each? • We can quantify the relative contributions of genetic and environmental variation to behavioral variation by thinking about heritability Heritability (H2) Heritability of colony size preference in the Lesser Kestrel One way to determine • Indicates what proportion of the total variance in a trait is heritability is to regress H2 = 0.53 due to variation in genes offspring phenotype with • Heritability can vary from 0.0-1.0 parental phenotype. • H2=0: Phenotypic variation not due to genotypic variation The slope of the regression 2 • H =1: All phenotypic variation due to genotypic variation line approximates – H2 is a relative measure, so depends on the population under heritability of trait (if avg. investigation trait value of parents used). – If H2 is high, can be impetus for investigations to identify the responsible gene(s) &/or the mechanisms of trait expression

(Serrano & Tella 2007)

Heritability (H2) Heritability (H2)

Total genotypic variation: VG = VA + VD + VI – Proportion of phenotypic variation in a population due to genetic variation where

VA = Additive effects, or the average effect of individual alleles VG on the phenotype 2 VD = Dominance effects, or the interaction between alleles at h = one locus VP VI = Epistasis, or the interaction between genes at different loci

• a measure of how strongly a behavioral phenotype is influenced by genetic differences between individuals 2 Heritability (H ) Heritability (H2)

Total phenotypic variation: VP=VG+VE+VI 2 H =VG/(VG+VE+VI) where: where:

VP= total phenotypic variation observed in a (behavioral) trait 2 H = heritability VG= variation in population due to genotype VG= variation in population due to genotype VE =variation in population due to environment VE =variation in population due to environment VI = variation in population due to interaction of VG with VE VI = variation in population due to interaction of VG with VE

• Gene-environment interaction (GEI) Rover and sitter foraging behavior in fruit flies – When environment has greater effect on one genotype than other • Research question: Do different behavioral polymorphisms in fruit flies exhibit gene- environment interactions? (Kent et al. 2009) Rover and sitter foraging behavior in fruit flies Rover and sitter foraging behavior in fruit flies

• Methods: – Exposed adult rover and sitter phenotypes to different levels of food availability (fed or food deprived) – Recorded movement from food patch – Used mass spectroscopy to determine compounds stored in head

Rover and sitter foraging behavior in fruit flies Rover and sitter foraging behavior in fruit flies • Results: – Fed rovers had much • Results: higher food-leaving – Rovers and sitters store scores than did food- food differently deprived rovers – No difference between • Conclusion: fed and food-deprived – Gene-environment sitters interactions affect both behavioral and metabolic traits

Figure 4.16. Adult rover and sitter food-leaving behavior. Mean (+ SE) food-leaving score for fed and food- deprived flies. Note the difference in the change in sitter (blue) and rover (orange) food-leaving scores between fed and food-deprived flies (Source: Kent et al. 2009). Development of Behavior Precocial young

Altricial young Behavioral Development

The changes in behavior and its underlying mechanisms in individuals from conception to death. The ontogenetic trajectory of a worker honey bee

This includes the behavior of individuals before they are born!

Waddington’s Epigenetic Landscape (1957) as a metaphor for Canalization: development a particular developmental process may follow a fairly fixed path, resulting in similarity between individuals of the same species.

When a ball moves to the left or right, the slope brings it back on course. Similarly, regulatory mechanisms keep development on track. Differentiation Humans pass the same developmental Certain internal and external events can trigger development milestones as they grow up. to change course. Different valleys represent the alternative phenotypic traits. - walk by 18 months - talk by 2 years of age - reach sexual maturity before the late teen years Y chromosome

These features would be described by Royal jelly Waddington as ‘canalised’.

Genetic, physiological and ecological factors are crucial for an individual’s life-history.

Some behavior patterns appear stable in a wide range of environments.

But these behavioral features can still be Spider’s web spinning modified by experience.

Rodent grooming

Infant smiling 6 Early social environment can affect behavioral development Blind babies start to smile at the same age as sighted babies. – Eg., hatching synchrony (23-24 days) in bobwhite quail chicks can be facilitated by chick clicks

But sighted people learn to modify their • clicks of more developed chicks hastened hatching date of less smiles according to their experience. developed chicks • clicks of less advanced chicks deferred hatching date of more developed chicks Blind people are less responsive and less expressive in facial expression, since they lack visual interaction.

Early social environment can affect behavioral development

– Female house mouse maturation & puberty ~35 days – if exposed to male urine or estrus female - earlier – if exposed to urine of group-caged females - later Social development early social environment can affect behavioral development What social problems confront an organism during its development?

1)Discriminating animate from inanimate objects 2)Identification & classification of prospective social partners 3)Attainment of skills necessary for competition & cooperation, establishing a social position Dispersers in social species must do this twice! 4)Acquire skills associated with mating: Identifying, finding, selecting, courting, and copulating 5)Develop parenting skills

Harlow Experiments Social deprivation: rhesus macaque studies

–Total Isolates: – required only 20 minutes/day of social contact! –non-normal behavior as juveniles – Rehabilitation: 6 month old isolate w/3 month –As adults- inappropriate social, sexual and old non-isolate parenting

–Surrogate mothers: –More normal self-directed behavior –Non-normal social and parenting

–Peer groups: –More normal behavior than when reared with mom alone Ballen et al. 2014

Early social environment can affect behavioral development – John Bowlby’s work on Attachment in Orphans from Imprinting WWII • often unable later to form any lasting, intense ‘love’ • Experience in early life relationships affects later social and • Critical importance of the young child’s relationship sexual preferences to at least one specific & dependable adult caregiver responsive to child’s emotional and physical needs • long-lasting emotional bonds that infants develop towards their primary caregiver Filial imprinting

•After hatching, chicks of many species approach & follow conspicuous objects to which they are exposed for a certain amount of time.

Filial Imprinting •Chicks will even form a social the process by which preference for inanimate objects. animals develop a social •Chicks become distressed when the attachment to a familiar object is removed. particular object

Critical (“sensitive”) Period for Filial Imprinting

* critical period

sensitive period *

mallard ducklings Colin Gregory