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Mendel’s seminal publication makes no mention of linkage, and many researchers have questioned whether he encountered linkage but chose not to publish those crosses out of concern that they would invalidate his independent assortment postulate. The garden pea has seven chromosomes, and some have suggested that his choice of seven characteristics was not a coincidence. However, even if the genes he examined were not located on separate chromosomes, it is possible that he simply did not observe linkage because of the extensive shuffling effects of recombination.


Mendel’s studies in pea plants implied that the sum of an individual’s phenotype was controlled by genes (or as he called them, unit factors), such that every characteristic was distinctly and completely controlled by a single gene. In fact, single observable characteristics are almost always under the influence of multiple genes (each with two or more alleles) acting in unison. For example, at least eight genes contribute to eye color in humans.

Concept in action

Eye color in humans is determined by multiple alleles. Use the Eye Color Calculator to predict the eye color of children from parental eye color.

In some cases, several genes can contribute to aspects of a common phenotype without their gene products ever directly interacting. In the case of organ development, for instance, genes may be expressed sequentially, with each gene adding to the complexity and specificity of the organ. Genes may function in complementary or synergistic fashions, such that two or more genes expressed simultaneously affect a phenotype. An apparent example of this occurs with human skin color, which appears to involve the action of at least three (and probably more) genes. Cases in which inheritance for a characteristic like skin color or human height depend on the combined effects of numerous genes are called polygenic inheritance.

Genes may also oppose each other, with one gene suppressing the expression of another. In epistasis    , the interaction between genes is antagonistic, such that one gene masks or interferes with the expression of another. “Epistasis” is a word composed of Greek roots meaning “standing upon.” The alleles that are being masked or silenced are said to be hypostatic to the epistatic alleles that are doing the masking. Often the biochemical basis of epistasis is a gene pathway in which expression of one gene is dependent on the function of a gene that precedes or follows it in the pathway.

An example of epistasis is pigmentation in mice. The wild-type coat color, agouti (AA) is dominant to solid-colored fur (aa). However, a separate gene C, when present as the recessive homozygote (cc), negates any expression of pigment from the A gene and results in an albino mouse ( [link] ). Therefore, the genotypes AAcc , Aacc , and aacc all produce the same albino phenotype. A cross between heterozygotes for both genes ( AaCc x AaCc ) would generate offspring with a phenotypic ratio of 9 agouti:3 black:4 albino ( [link] ). In this case, the C gene is epistatic to the A gene.

A cross between two agouti mice with the heterozygous genotype AaCc is shown. Each mouse produces four different kinds of gametes (AC, aC, Ac, and ac). A 4 × 4 Punnett square is used to determine the genotypic ratio of the offspring. The phenotypic ratio is 9/16 agouti, 3/16 black, and 4/16 white.
In this example of epistasis, one gene ( C ) masks the expression of another ( A ) for coat color. When the C allele is present, coat color is expressed; when it is absent ( cc ), no coat color is expressed. Coat color depends on the A gene, which shows dominance, with the recessive homozygote showing a different phenotype than the heterozygote or dominant homozygote.

Section summary

Alleles do not always behave in dominant and recessive patterns. Incomplete dominance describes situations in which the heterozygote exhibits a phenotype that is intermediate between the homozygous phenotypes. Codominance describes the simultaneous expression of both of the alleles in the heterozygote. Although diploid organisms can only have two alleles for any given gene, it is common for more than two alleles for a gene to exist in a population. In humans, as in many animals and some plants, females have two X chromosomes and males have one X and one Y chromosome. Genes that are present on the X but not the Y chromosome are said to be X-linked, such that males only inherit one allele for the gene, and females inherit two.

According to Mendel’s law of independent assortment, genes sort independently of each other into gametes during meiosis. This occurs because chromosomes, on which the genes reside, assort independently during meiosis and crossovers cause most genes on the same chromosomes to also behave independently. When genes are located in close proximity on the same chromosome, their alleles tend to be inherited together. This results in offspring ratios that violate Mendel's law of independent assortment. However, recombination serves to exchange genetic material on homologous chromosomes such that maternal and paternal alleles may be recombined on the same chromosome. This is why alleles on a given chromosome are not always inherited together. Recombination is a random event occurring anywhere on a chromosome. Therefore, genes that are far apart on the same chromosome are likely to still assort independently because of recombination events that occurred in the intervening chromosomal space.

Whether or not they are sorting independently, genes may interact at the level of gene products, such that the expression of an allele for one gene masks or modifies the expression of an allele for a different gene. This is called epistasis.

Art connections

[link] What ratio of offspring would result from a cross between a white-eyed male and a female that is heterozygous for red eye color?

[link] Half of the female offspring would be heterozygous (X W X w ) with red eyes, and half would be homozygous recessive (X w X w ) with white eyes. Half of the male offspring would be hemizygous dominant (X W Y) with red eyes, and half would be hemizygous recessive (X w Y) with white eyes.

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Questions & Answers

must all prokaryotic cells posses a cell wall?
chris Reply
what is biology?
Cathy Reply
biology is basically the study of life
that's true
it's the study of living things
What is a celiac disease
Falase Reply
distinguish between properties and characteristics
Elee Reply
what are organelles
organelles are substances that makes up a cell
what is biology
Prevail Reply
biology is science that studies life
it's the study of living and non living things
now that we have an estimate for the diameter of the cell.what estimate can we make about the volume of the cell?
faxhood Reply
why too much insulin result in low blood sugar
Leri Reply
For example, too much exercise can cost you to lose to much weight. Too much insulin will pull to much sugar out of your systemic system into your cells.
difference between DNA and RNA
Eyitayo Reply
DNA is deoxyribonuclaic acid. Deoxy refers to a lack of oxygen. The Ribose moity is missing an OH group. I think it is missing from the second C of the ring. RNA is ribonucleic acid. DNA has our genetic code in on it. RNA is translated from DNA and carries the blue print for protein synthesis.
The OH group on RNA prevents it from being reactive. But it is very unstable though. Would you want such a power tool floating around in your body, no. And you have three types of RNA: mRNA, tRNA and rRNA. Please let me know it this helped?😄
Eric answered this question perfectly
what determines the aeration level in the soil
Shola Reply
what is homeostasis?
Sarita Reply
It means balance in a biological system.
What is biology
Don Reply
Biology z the study of life
what's biology
biology is the study of living nd none living organism
Biology is the study of life
yes Sir
what's cell biology
biology is the study of life
Biology is a science subject that deals with the study of living things and how they interact with there environment
what is asexual reproduction,?
Awoi Reply
A type of reproduction which does not involve the fusion of gametes or a change in the number of chromosomes
Reproduction without sex... In which form a single organism or cell makes a copy of itself.
Please explain the concept of mitosis and meiosis
I guess you could use it for study buddies and brushing up on what you need to
what is mitosis
Asexual reproduction?
why pepsin and trypsin released in active form?
mitosis is the type cell division in which two daughter cells have same no. of chormosomes
chromosome number remains the same in mitosis
Yrr help me.
Physical chemistry..... Koi h jo mujhe physical chem ki notes send kr ske
what is asexual reproduction
what makes golgi body in plants
Abdulkareem Reply
name the membrane of the plants
how can turners syndrome be corrected before birth
which animal survive from being preyed just because of being humble, slow, and not aggressive
Plants have golgi body's also. Plants are eukaryotic cells. And membrane bound organelles are a characteristic of eukaryotic cells. Moreover golgi body's are creatted from the ER. Also do not forget plants have plastids and animal cells do not.
During organs transplantation, the organs cannot be taken from just anybody since the graft would be rejected sooner or later due to
Liter Reply
Non-MHC compatibility on the organ and an attack from the patient's immune system.
what makes golgi body in plants
why trypsin and pepsin released in active form
Let us remember MHC'S on our cells. This is how our cells determine self from n o n s e l f. Transplanted tissue has to have a certain amount markers. These have to match to the recipiant's markers. Even with this, immunosuppresant medacine is prescribed to the recipient.
Even with these measures the body may still reject the transport. This can occur even after the recipient excepting the transport for some time.
what is integument system
Joy Reply
This system is our skin. This includes the skin lining our alimentary system which includes the tissue from our mouth to our anus. Our skin is our largest organ system. It is mostly made up of epithelial tissue.

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Source:  OpenStax, Concepts of biology. OpenStax CNX. Feb 29, 2016 Download for free at http://cnx.org/content/col11487/1.9
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