WAP 214 PRINCIPLES OF ANIMAL BREEDING Office hours: 0900hrs-1645hrs (Mon-Fri) Textbooks: Richard M. Bourdon. 1997. Understanding animal breeding,Dr S.M. Makuza.Animal Breeding.ZOU Module. Prerequsite:AGRO101 -Principles of Genetics COURSE OBJECTIVES: The course is designed to enhance your understanding of genetic principles and their applications on genetic improvement of livestock populations. During the first part of the course we will review concepts of classical (Mendelian) inheritance and population genetics. Then, we’ll learn how to calculate and interpret basic statistics used to describe a given population and on genetic evaluation. We’ll then discuss selection principles for short and long term responses on herd improvement. We’ll focus on interpretation of performance records and estimated genetic values, including information produced by national genetic evaluation programs, such as breed association sire summaries. STUDENT OUTCOMES: After attending the course, the students will able to: 1. Use statistics to describe the genetic and phenotypic variation of a given population. 2. Explain the use of phenotypic information from the animal and its relatives in estimating transmitting ability. 3. Understand the information generated on a genetic evaluation program and how to apply this information on animal breeding. 4. Understand concepts of breeding goal, selection index, heritability, repeatability, correlation, accuracy, selection intensity, generation interval and their effects in making genetic change. 5. Relate the Hardy-Weinberg Law to forces that causes changes on genetic and genotypic frequencies. 6. Design breeding programs using the concepts listed above to achieve defined breeding goals. 7. Calculate relationship coefficient and inbreeding and understand its effects at the application level. 8. Infer relationships between reproductive rate, longevity on selection intensity and generation interval. 9. Characterize heterozygosis and heterosis and how to explore it through systematic crossbreeding 10. Understand the role of molecular genetics and new technologies on genetic improvement. 11. Understand the biological and biochemical basis for genetic variability and heritability 12. Understand advantages and limitation of multiple trait selection. 13. Explain the impact of new technologies on livestock improvement. GRADING:Coursework-30% 2 assignments (10%)+ 1 mid-semester test (20%)exam-70% Final COURSE OUTLINE: I. Introduction, History of Animal Breeding & Genetics (Chapters 1 and 2)-Description of various domestic livestock species, breeds, their origin, productive & adaptive characteristics. II. Review of Basic Principles of Mendelian Inheritance 1. Cellular aspects of genetics 2. Mendel's laws of Inheritance. 3. Functional aspects of Dominance and Co-dominance (Gene Action). 4. Testing for genetic abnormalities, Pleiotropic effects, variable expressivity,Incomplete Penetrance. III. Population Genetics 1. Gene and genotypic frequencies 2. Hardy-Weinberg Law 3. Forces that change gene frequency a) Mutation b) Migration c) Selection d) Random drift e) Mating Systems The importance of changing gene/genotype frequencies in livestock improvement. IV. Principles of selection-influence of selection, selection response, 1.Basic statistics for animal breeding a) Mean b) Variance and standard deviation c) Regression and correlation d) Normal distribution, statistical significance and confidence limits. 2. Relationships and inbreeding. 3.Quantitative traits-the quantitaive traits model, phenotypic and genetic variation in animals (causes and estimation). Genotype value, breeding value, additive genetic value, dominace genetic value, epistatic genetic value. 4.Genetic parameters- Heritability and repeatability V.Heredity and environment-Genotype*Environmental interaction,environmental effects, maternal effects, phenotypic corrrelation,environmental corrrelations, genetic correlations. Cattle breeds in Zimbabwe Bos taurus Bos indicus -polled/short horned -hairy and rough, diificult control of external parasites. -insignificant humps The above are the phyical traits, below are the production traits. -low disease resistance in Zim. -Highly selective-demand good quality feed.Not browsers. -low meat to bone ratio. -low fertility in Zim. Examples of breeds -Charolais, Sussex, Simmental (Dual purpose). -Hereford, Limousine, Gelbviah,Belgian blue, Aberdeen Angus, Blonde Aquatine, Marchigiana (Beef). -Composite breeds-Beefmaster, Bonsmara,Santa Gertrudis, Charbrays, Simbras,Brahford. -Dairy breeds-Jersey, Red Dane, Holstein, Aryshire, Guernsey, Dexter, Milk Shorthorn. Students to have an appreciation of the physical and production characteristics of each breed. -big horned -smooth shiny coats -pronounced humps. -high disease resistance. -non-selective, good browsers, survive in drought periods. - high meat to bone ratio. - average fertility. -Brahaman (white/red), Tuli (9 colors), Nguni (red/back with white spots),Mashona (red/black), Afrikander(big huge horns),Boran (like the Brahman but short legs-fron Kenya prevalent in the Mvurwi area.), Sahiwal (kenya), Angoni (Zambia very short and very small150kg). Composite Breeds. Defn: A breed made up of two / more component breeds and designed to benefit from hybrid vigour without crossing with other breeds. Examples, 1. Beefmaster-Brahman*Hereford=F1*Dairy shorthorn 2. Bonsmara-Afrikander*Hereford=F1*Dairy shorthorn. 3. Santa Gertrudis-Brahman*Sussex. Students to explore other different types of livestock and their breeds, ( sheep, goats, poultry, pigs). Review of basic principles Of Mendelian Inheritance. Key words; Gene, DNA, chromosome, locus, allele, genotype, honozygous, heterozygous, segregation, germ cell, meiosis, independent assortment, linkage, crossing over, Punnet square, Mendelian Sampling,Dominance, no dominance, partial dominance, complete dominance, overdominance, epistasis. Defination of terms: Gene-basic unit of inheritance, can also be described as a segment of DNA at a specific location on a chromosome. DNA- a complex molecule that forms the genetic code. Chromosome- a strand of DNA and associated proteins present in the nucleus of every cell of a living organism. Locus- the specific location of a gene on a chromosome. Plural loci.They are two genes at each locus, one partenaland one martenal. Allele- an alternative form of a gene. Genotype- a combination of genes at a single locus or a number of locus. Homozygous Genotype- a one locus genotype consisting of functionally identical genes. Heterozygous Genotype-a one locus genotype consisting of functionally different genes. Segregation- the separation of paired genes during germ cell formation. Germ cell- sex cell (sperm/egg). Meiosis- process of germ cell formation.P Punnet Square- a two dimensioal grid used to determine the possibe zygotes obtainable from a mating. Mendelian Sampling-the random sampling of parental genes caused by segregation and independent assortment of genes during germ cell formation and by random selection of gametes in the formation of the embryo. Dominance- an interaction of genes at a single locus such that in the heterozygous one allele has more effect than the other. The one that is masked is said to be recessive. Complete Dominance- A form of dominace in which the expression of the heterozygote is idebtical to the expression of the homozygous dominat genotype. Partial Domiance-A form of dominace in which the expression of the heterozygote is intermediate to the expression of the homozygous genotypes and closely resembles the homozygous dominant genotype (which can be recessive as well). No dominace-A form of dominance in which the expression of the heterozygote s exactly midway between the expression of the homozygous genotypes. Overdominance- a form of dominace in which the expression of the heterozygote is outside the range defined by the expressions of the homozygous genotypes and most closely resembles the expressions of the dominant genotype. Epistatis- An interaction among genes at different loci such that the expression of genes at one locus depends on the alleles present at one or more loci. Pleiotropy-a genetic situation in which one gene /sing;e allele affects more than one distinct qualitative/ quantative trait of an individual animal. Variabile expressivity- similar individuals genetically either show up/ are supressed due to environmental variations. Incomplete penetrance-the genotype of a trait is not expresssed in the phenotype at all. Mendel's Laws of Inheritance. 1. Law of Segreagation-In the formation of a germ cell (meiosis) the two genes at a locus in the parent cell are separated, only one gene being incorporated into each germ cell. 2. Law of Independent Assortment-genes assort independently during meiosis if all possible gametes are formed in equal proportions. (A given gene from one locus must have an equal probability of being present in the same germ cell with either of the two genes from some other locus).Segregation at one locus does not influence segreagation at another. Assupmtions of the law: 1. There is no linkage i.e. The loci dnt occur on the same chromosome if they do then some genes will occur at a higher frequencies than others. 2. There is no crossing over i.e no exhange of chromosomal segemts between homologs, if it happens then the genes will be rearranged and there will be differences in their frequencies. 3. Some genes are lethal, some genes are affected by the environment (phenocopies), Illustrations of Mendel's laws: Parental Genotype: JJBB JJBb JjBb Possible gametes: JB JB Jb JB Jb jB jb Example above show 2 locus genotype (with homozygous and heterozygous genotyepes). For law 1, each gamete contains only one gene from each locus. NB: the original 2 locus genotype contains 4 genes altogether, the resultant germ cell contains, as a Rule, germ cells contain half the number of chromosomes and therefore half the number of genes of normal body cells. For law 2, the homozygous genotype (JJBB) can produce 1 gamete, , the partial heterozyguos can produce two and the completely heterozygous can produce 4 gametes. For the latter if all four gametes occur in equal proportions then the genes have assorted independently unless otherwise there is linkage and crossing over. Gamete Selection: The Punnet square When two individuals mate all possible gametes have an equal chance of contributing to a zygote. This is a random process. The Punnet square is a commonly used device for determining the possible zygotes from the mating of two parental genotypes. f JB Jb jB jb JJBB------1 JJBb----------2 JjBB--------3 JjBb----------4 JB horizontal: female, vertical male (punnet square showing the possible gametes from mating two animals of the same 2 locus genotype JjBb). There are 9 possible unique genotypes. It is possible therefore to determine the likelihood of any particular offspring genotype, and if you know the phenotype associated with each genotype (for simply inhertaed traits not plygenic traits) u can also determine the expected proportions of offspring phenotypes. A gud example is coat color in Shorthorn cattle ( red, white or roan).The colors are controledd by the white locus RR=red, Rr= roan, rr=white. Using the Punnet square u can mate 2 roans (illustrate), the results show three offspring genotypes and phenotypes in the ratio 1:2:1 (red, roan and white).NB this is an expectation. We cannot control the outcome of matings as there is random sampling of parental genes and random selection of gametes during the formation of an embryo (Mendelian Sampling). This leads to genetiv variation through segreagation and recombination. Number of unique gametes an indv can produce=2 raised to the power n, where n is the number of loci at which the individual is heterozygous. (some examples- how many gametes can be produced by an indv, with the foll genotype AaBbCc Ans=8) show working. If the indv (AaBBCcDdee) is mated to a female with the following genotype AABbCcDdEe, the number of unique zygotes = 3^n* 2^m where n is the number of loci at which both parents are heterozygous and m is the number of loci at which one parent is heterozygous. How many unique gametes can the sire produce=Ans (8) How many unique gametes can the dam produce=Ans (16) How many unique gametes can be produced from this mating=Ans( 72). show working. Examples of diffeernt types of Gene Expressions (Gene Action) Complete domiance- common in a number of simply inherited traits e.g polledness P (polledness) is dominant over p (horned). Traits have more than one genotype for a phenotype ( Polled cattle can be PP/Pp. e.g purebred Angus (PP&BB -black) * purebred Hereford ( horned&bb). F1 is heterozyguos polled black animals. Show Punnet square. Crosing the F1 gen, results in 39polled (black) and one horned (red)., 3 black horned, 3 polled red.show working. J'J' JJ JJ' Partial domiance- hyperkalemic periodic paraylsis HYPP in horses. Causes muscle tremors resulting in heavy shaking and collapse in horses but also leads to heavy muscling in horess necessary for increased speed in horses hence it has peroetuated coz Breeders unknowingly select for it. J'J' JJ' JJ JJ' is more closer to JJ hence J is the dominant allele. b) J'J' JJ' JJ in this case J' is the domiant allele as exemplified by the HYPP syndrome in horses. Another xample is the bull dog (short legs and compavct bodies) condition in Dexter cattle (the trait is dominant over the normal).Partail domiance can also be used to explain the phenomenon of additive gene actio and non-additive gene action.This is for the former two alleles may be equal in power so that the phenotype of the heterozygote is midway between trhe homozygotes.It is used with ref to quantitative traits;milk production, weaning weight and backfat thickness in pigs.In non-additive gene action the heterozygot does not lie between the homozygotes. No dominance- resistance to tuberculosis, res gene (T^r) animals survive 100% of the tym, susceptible gene animals survive 40% of the tym. Heterozygous gene animals survive 70% of the tym hence there is no dominace in the locus. JJ JJ' JJ Overdominance- survivabiltiy of wild rats, both homozygoytes not affected by the poison warfarin, rats without the resistant gene succumb to warfarin, the warfarin gene displays overdominance. The phenomenon can be expressed in 2 ways, a) b) J'J' JJ' JJ J'J' JJ' JJ Epistatis (non-additive gene action) Epistasis affects the kinds and proportion of phenotypes we can expect from matingsin simply inherited traits.It is also a source of hybrid vigour and inbreeding depression in polygenic traits. Examples of epistasis, explain, the case of an albino (recessive epistasis) where color is infuence d at the B-extension locus. Dominace epistasis-coat color in sheep, white dominant. Apllication of Gene Action in Livestock Improvement Dominace,Epistasis are implicated in theories which explain hybrid/heterosis. Additive gene action is exploited by breeders in selection and culling programmes. Testing for careers with recesive alleles. This is done so as to eliminate the recessive genes which are usually lethal.3 methods are used. Mating the suspect male to the known heterozygos female. If the male is a career, ¼ of the f1 will be recessive and exhibit the condition. Mating a sire to its own daughters.The expected frequency is 7 heterozygos and 1 defective.the test requires 17 progeny to have a 10% chance of success. Mating with affected females (homozygos), 1 affected offspring show that the male is a career. Pleiotropic Effects Usually common in traits which are postively corelated e.g. Weaning weight and birth weight they have genes whic are common. Also they are associated with the lethal effects of the bull dog condition in Dexter cattle.The product of a single gene will affect more than one trait. Variable Expressivity A G*E interaction. e.gs include mulefootedness in pigs (dominant) and cattle (recessive), Porcine Stress Syndrome (PSS)- affects well muscled pigs (labored breathing which can be fatal). Incomplete Penetrance The animal appears normal though it is affected, e.g polydactylty in fowl and humans. The indv has an extra digit on the foot/finger.The gene for plydactylty is domiant over that of the normal but fails to express iself in the phenotype. PRINCIPLES OF POPULATION GENETICS The goal of animal breeding is to change populations and not individuals therefore we are concerned about programms that have have the capacity to change gene and genotype frequencies in populations. Keywords and concepts: Gene and genotype frequency, H-W law, mutation, migration, selection, mating systems, inbreeding, outbreeding, relationships, Defiantio of terms: Population genetics- the study of factors affecting gene and genotype frequencies in a population. (usually limited to simply -inherited traits). Gene/allelic frequency- the relative frequency of a particular allele in a population. How common is the allele relative to other alleles in a population.. Genotypic frequency- relative frequency of a particular one locus genotype in a population. Mating System- is a set of rules for mating ( Inbreeding, random mating,assortative mating, outbreeding). Relationship-full-sib (full brothers/sisters), half sibs, parent-offspring. Hardy-Weinberg law- a state of constant gene and genotype frequencies occuring in a population in the absence of forces that bchange those frequencies. Mutation- the process that alters DNA to create new alleles. Migration- the movement of indv into or out of a population.Se Selection- the process that determines which individuals become parents, how many offspring they may produce, and how long they remain in the breeding population. Inbreding-the mating of relatives. Outbreeding-the mating of unrelated individuals. Relative frequency-ranges from 0-1. The frequency of the dominant allele is represented by the lowercase letter p. The frequency of the recessive allele is represented by the lowercase letter q. Homozygous dominant=P heterozygous =H homozygous recessive=Q P=p^2 H=2pq Q=q^2. Number of p genes=P+1/2H number of q genes= Q+1/2H p+q=1 Calculations A flock of 100 Andalusians consists of 36 black (BB), 44 blue (Bb) and 20 white (bb). Calucate the gene and genotype frequencies. Total number of genes=100*2 genes for each individual i.e 200. p= 36+1/2(44)=58*2=116 black genes. q=20+1/2(44)=42*2=84 white genes. f(p)= 116/200=0.58 f(q)=84/200=0.42 f(P)=36/100=0.36 F(Q)= 20/100=0.2 f(H)= 44/100=0.44 NB: P+Q+H=1 FACTORS THAT AFFECT GENE AND GENOTYPE FREQUENCIES IN A POPULATION. These apply to both quantative (polygenic) and qualitative traits (simply-inherited). Examples of Qualitative traits-affected by few loci. Polledness and coat color. Examples of Quantative traits-affected by many loci. Growth rate, carcasse composition, milk yield, weaning weight, backfat thickness, egg production, feed efficiency, yearling weight, fat percentage , protein yield. 1.Selection- Selection increases the gene frequency (immediate action-genotype fequencies take time )of favorable alleles in a population.We select for animals with a high breeding value ( better sets of genes) so that the next gen may have an improved breeding value. Show this on a diagram.Read more about selectio-natural versus artifial selection ( replacement selection, culling, phenotypic selection-for those traits that are highly heritable, pedigree and progeny selection, collateral selection-use of contemporaries, performance testing-use of sire summaries). 2. Mating Systems ( Random mating, assortative mating, outbreeding and inbreeding-explain this)- this has an effect when combined with selection.The more common use of mating systems is to change genotypic frequencies by increasing the number of homozygous /heterozygous gene combinations.Gene frequencies may/ may not change as a result. Breeders use mating systems to achieve the following-- a) to produce offspring with extreme breeding value to increase rate of genetic change, b) to make use of complementarity c) to obtain hybrid vigour. We have two catergories for the purposes of this discussion: Inbreeding and Oubreeding. Inbreeding – increases the frequency of homozygous genotypes. Types of relationships: Full sib, half sib, common ancestor (pedigree). Show relationship diagram and explain inbreeding.Inbreeding can be used to create the breeds within species or lines within breeds that when crossed produce hybrid vigour. Oubreeding (Crossbreeding)- increases hybrid vigour by ncreasing heterozygosity.Example there are two unrelated populations. P1=0.8 , q1=0.2 & p2=0.1 q2=0.9 the differences in the frequencies reveal that indeed the populations are unrelated. Crosing the two populations (Bb* Bb) will result in F1, BB= 0.08 Bb=0.74 bb =0.18 (from punnet square). NB: The increase in heterozygosity and the decrease in homozygousity. f(p)=( P+ 1/2H)*2 divided by the total number of genes= 0.45 f(q)=0.55. Mating F1, will result in BB=0.2025 Bb=0.495 bb=0.3025 NB; An increase in homozygosity as this is now a form of inbreeding. However on calcultion of the gene frequencies, they dnt change p=0.45 and q=0.55. i.e pF2=Pf2+1/2Hf2 =0.45 and qf2=Qf2+1/2Hf2=0.55. The chi-square goodnes of fit test is used to test whether a population is in H-W equilbrium. This phenomenon is known as the H-W law of equilbrium (1908)The frequencies will only change under conditions of selection, mutation and migration. Futher Assumptions of the law. 1. Large populations. 2. Migrations can be negligible. 3. Random mating-only one generation is needed to reach equilibrium. 4. There is no selection. 5. There is no mutation. 6. There is normal gene segregation. 7. There is equal fertility of parents anf equal ferilising capacity of gametes and reproduction is sexual 8. There is equal viability and equal gene frequencies in male and female parents. 9. The generations are non overlapping. 10. The gene frequency is the same in parents and offspring. If gene frequencies are the sane in both sexes a population reaches H-W after one generation, if not H-W is reached after two generations.With sex-linked loci this may take several generations. Why? Example- In a population of 1000 Hampshire pigs 910 are belted and 90 are solid colored. What is the frequency of the genotypes. Ans F(recesive)=90/1000 Q=q^2 Q= 0.09 hence q=square root of Q= 0.3 and therefore p=0.7 p^2=P P=0.49 H=2pq= 0.42 number of heterozygosites= 0.42*1000=420 numbr of homozygosites dominant=0.49*1000=490. Note that this is a case of coimplete domiance. 3) Mutation- explain how it changes genotype frequencies.Use examples. -Mutatios are very slow. -Mutations may occur in any direction, the major role is to present new genetic material(desirable/undesirable) into the population.Selection then increases the frequency of the desirable mutation e.g polledness. -Mutation has a minimum impact on quantative traits aand is not useful in domestic animals. 4) Migrations-use examples to explain.(imports of exotic breeds). This can be a rapid way of changing gene frequency, the rate of change will howver depend on the following. Frequencies of the gene (s) in the immigrants must be high i.e. The differences btwn imm and natives. The number of immigrants introduced in2 the native population. 5) Random Genetic Drift (Effect of chance) This is a chance change in gene frequency outside any influence, it is most likely in small popualtions.This is largely due to chance sampling of parents which affects the offspring.e.g the appearance of horned cattle in a herd selected for polledness.horned Holstein cattle in Zim. The importance of changing gene and genotype frequencies in livestock 1. Selection increases gene frequency and in turn incrreases performance. 2. Changing gene frequency is the only way to make a long term improvement e.g im milk production, eggs, higher grwoth rates. 3. Changing gene frequency is a permanent change buit we cannot change gene action. 4. Long term change in gene action is the central issue in selection. 5. Mating systems can change genotypic frequencies but one can lose wht he/she has gained.