Non-Mendelian Genetics (College Board AP® Biology): Study Guide
Gene linkage
Some patterns of inheritance do not follow Mendel's laws
Their observed phenotypic ratios among the offspring differ significantly from the predicted ratios
Patterns of inheritance of traits that do not follow ratios predicted by Mendel’s laws and can be identified by quantitative analysis
One example is gene linkage where some genes are not inherited independently from one another
Autosomal linkage
Loci (singular: locus) refers to the specific linear positions on the chromosome that genes occupy
Linked genes located on chromosomes 1-22 (not sex chromosomes) are called autosomes and are examples of autosomal linkage
Dihybrid crosses and their predictions rely on the assumption that the genes being investigated behave independently of one another during meiosis
However, not all genes assort independently during meiosis
Some genes which are located on the same chromosome display autosomal linkage and stay together in the original parental combination
Linkage between genes affects how parental alleles are passed onto offspring through the gametes
The distance between linked genes on a chromosome can be mapped using the probability that the linked genes will be inherited together
Identifying autosomal linkage from phenotypic ratios
In the following theoretical example, a dihybrid cross is used to predict the inheritance of two different characteristics in a species of newt
The genes are for tail length and scale color
The gene for tail length has two alleles:
Dominant allele T produces a normal length tail
Recessive allele t produces a shorter length tail
The gene for scale color has two alleles:
Dominant allele G produces green scales
Recessive allele g produces white scales
Without linkage
Normal Mendelian ratios would result if there is no linkage
The outcomes for this dihybrid cross if the genes are unlinked are as follows

Predicted ratio of phenotypes in offspring =
1 normal tail, green scales : 1 normal tail, white scales : 1 short tail, green scales : 1 short tail, white scales
Predicted ratio of genotypes in offspring =
1 TtGg : 1 Ttgg : 1 ttGg : 1 ttgg
With linkage
However, if the same dihybrid cross is carried out but this time the genes are linked, we get a different phenotypic ratio
There would be a 1 : 1 phenotypic ratio (1 normal tail, green scales : 1 short tail, white scales)
This change in the phenotypic ratio occurs because the genes are located on the same chromosome
The unexpected phenotypic ratio, therefore, shows us that the genes are linked
Parental phenotypes: normal tail, green scales x short tail, white scales
Parental genotypes: TG tg tg tg
Parental gametes: (TG) or (tg) (tg)

Predicted ratio of genotypes in offspring =
1 (TG)(tg) : 1 (tg)(tg)
Predicted ratio of phenotypes in offspring =
1 normal tail, green scales : 1 short tail, white scales
Sex linkage
Some genetic diseases in humans are sex-linked
Inheritance of these diseases is different in males and females
Sex-linked genes are only present on one sex chromosome and not the other
This means the sex of an individual affects what alleles they pass on to their offspring through their gametes
If the gene is on the X chromosome males (XY) will only have one copy of the gene, whereas females (XX) will have two
This occurs in mammals and flies
If the gene is on the X chromosome, males (XY) will only have one copy of the gene, whereas females (XX) will have two
There are three phenotypes for females:
normal
carrier
has the disease,
Males have only two phenotypes
normal
has the disease
In certain other species, the chromosomal basis of sex determination is not based on X and Y chromosomes
This occurs in birds which have sex chromosomes called Z and W; these determine the sex of the bird
Bees have a genetic system called haplodiploidy that determines the sex of offspring; female bees are diploid and develop from fertilized eggs, while males are haploid and develop from unfertilized eggs
Examiner Tips and Tricks
The expected notation when writing about sex linked alleles is to use upper case 'X' and 'Y' for the chromosome, next to superscript letters to represent the allele. For example
XfXf Homozygous female who has hemophilia
XFXf Heterozygous female who is a carrier
XfY Male who has hemophilia
Worked Example
The genetic diagram below shows how two parents with normal factor VIII can have offspring with hemophilia.
What is the predicted ratio of genotypes in the offspring?
Parental phenotypes: carrier female x normal male
Parental genotypes: XFXf XFY
Parental gametes: XF or Xf XF or Y

Answer:
1 female with normal blood clotting : 1 carrier female : 1 male with hemophilia : 1 male with normal blood clotting
Predicted ratio of genotypes in offspring: 1 XFXF : 1 XFXf : 1 XFY : 1 XfY
Examiner Tips and Tricks
Make sure to include all of your working out when constructing genetic diagrams. It is not enough just to complete a Punnett grid, you need to show that you have thought about the possible gametes each parent can produce. Also, remember to state the phenotype as well as the genotype of the offspring that result from the cross.
Read the questions carefully when answering sex-linked inheritance questions – is the question asking for a probability for all children or is it asking about specific sex (males or females)?
Multiple genes
Multiple genes and physiological processes determine many traits and therefore do not follow Mendelian patterns
Traits from multiple genes do not follow predictable ratios like 3:1 or 9:3:3:1
Examples of such traits include height, intelligence and skin color, and result from multiple genes working together
These are called polygenic traits and show a range of phenotypes (continuous variation), rather than simple dominant or recessive patterns
Genes can also interact with each other; one gene affects or masks the expression of another, influencing the phenotype in complex ways: this is called epistasis
The influence of the environment affects the expression of traits due to the interaction with genes which may further complicate inheritance patterns
Non-nuclear inheritance
Not all characteristics are determined by genes carried on chromosomes in the nucleus in eukaryotes
Some genes are located within organelles elsewhere in the cell, away from the nucleus
Mitochondria, chloroplasts (and other plastids) carry DNA
This non-nuclear DNA is thought to date back to the origins of eukaryotic life and gives further supporting evidence to the theory of endosymbiosis
This DNA is not passed onto future generations in the same ways as nuclear DNA, so the traits it carries are inherited in non-Mendelian ways
The genes they carry are randomly assorted to daughter cells
In animals, mitochondria are transmitted by the female egg cell and not by sperm cells
Therefore, characteristics coded for in the mitochondrial DNA are inherited through the maternal line
Similarly, in plants, mitochondria and chloroplasts are passed on in the ovule and not by the pollen grains
Therefore, characteristics coded for in the mitochondrial and chloroplast DNA are inherited through the maternal line in plants also
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