Linkage and Crossing over
Linkage and crossing over explain why some genes do not assort independently. This page first covers why Mendel's work stayed unrecognised until 1900, the discovery of chromosomes, and how Sutton and Boveri used meiosis to explain Mendel's laws in the chromosomal theory of inheritance. It then covers Morgan's work on Drosophila, linkage, recombination and Sturtevant's gene maps, as in the NCERT Class 12 chapter Principles of Inheritance and Variation. NEET often asks Morgan's recombination values and the meaning of tight and loose linkage and crossing over.
- Mendel published his work in 1865; it stayed unrecognised till 1900.
- ★ Must learn 1900: de Vries, Correns and von Tschermak independently rediscovered Mendel's results.
- ★ Must learn Sutton and Boveri: chromosomes behave like genes; Sutton called the synthesis the chromosomal theory of inheritance.
- Chromosomes and genes both occur in pairs, segregate at gamete formation and assort independently.
- Morgan verified the theory using Drosophila melanogaster: life cycle about two weeks, many progeny, easily told sexes.
- ★ Must learn Linkage: physical association of genes on a chromosome. Recombination: generation of non-parental gene combinations.
- ★ Must learn White-yellow: 1.3% recombination (tight linkage). White-miniature wing: 37.2% (loose linkage).
- ★ Must learn Alfred Sturtevant used recombination frequency as a measure of gene distance to map genes.
- Genetic maps are the starting point for sequencing whole genomes, as in the Human Genome Project.
1. Why Mendel's Work Stayed Unrecognised
- ★ Exam imp Mendel published his work on the inheritance of characters in 1865, but it remained unrecognised till 1900.
- Communication was not easy in those days, so his work could not be widely publicised.
- His concept of genes (factors) as stable, discrete units that did not blend was not accepted by his contemporaries as an explanation for the apparently continuous variation seen in nature.
- His approach of using mathematics to explain biological phenomena was totally new and unacceptable to many biologists of his time.
- Although his work suggested that factors were discrete units, he could not give any physical proof that factors existed or say what they were made of.
2. Rediscovery of Mendel and the Discovery of Chromosomes
- ★ Exam imp In 1900, three scientists, de Vries, Correns and von Tschermak, independently rediscovered Mendel's results on the inheritance of characters.
- By then, advances in microscopy allowed scientists to observe cell division carefully.
- This led to the discovery of structures in the nucleus that appeared to double and divide just before each cell division.
- These were called chromosomes (coloured bodies), as they were seen by staining.
- By 1902, the movement of chromosomes during meiosis had been worked out.
3. The Chromosomal Theory of Inheritance
- ★ Exam imp Walter Sutton and Theodore Boveri noted that the behaviour of chromosomes was parallel to the behaviour of genes.
- They used chromosome movement to explain Mendel's laws.
- Recall: chromosomes behave differently in mitosis (equational division) and meiosis (reduction division).
- Chromosomes as well as genes occur in pairs.
- The two alleles of a gene pair are located at homologous sites on homologous chromosomes.
A comparison between the behaviour of chromosomes and genes
| Chromosomes | Genes |
|---|---|
| Occur in pairs | Occur in pairs |
| Segregate at the time of gamete formation, so that only one of each pair passes to a gamete | Segregate at gamete formation, and only one of each pair passes to a gamete |
| Independent pairs segregate independently of each other | One pair segregates independently of another pair |
3.1 Independent alignment of chromosome pairs
- In meiosis I, the two chromosome pairs can align at the metaphase plate independently of each other, and so separate independently at anaphase.
- Compare the four coloured chromosomes below: long orange and long yellow form one pair; short green and short red form the other.
- Possibility I: the long orange and short green chromosomes go to the same pole, and the long yellow and short red to the other.
- Possibility II: the long orange chromosome goes with the short red, and the long yellow with the short green.
- Both are equally likely, so four kinds of germ cells form in equal numbers.
- Sutton and Boveri argued that the pairing and separation of a pair of chromosomes would lead to the segregation of the pair of factors they carried.
- ★ Exam imp Sutton united the knowledge of chromosomal segregation with Mendelian principles and called it the chromosomal theory of inheritance.
Which three scientists rediscovered Mendel's work in 1900?
Who proposed the chromosomal theory of inheritance?
Where are the two alleles of a gene pair located?
In a table comparing chromosomes and genes, how can you tell which column describes chromosomes?
4. Morgan and the Fruit Fly
- The chromosomal theory was verified experimentally by Thomas Hunt Morgan and his colleagues.
- This also revealed the basis for the variation produced by sexual reproduction.
- ★ Exam imp Morgan worked with the tiny fruit fly, Drosophila melanogaster, which was very suitable for such studies.
Why Drosophila suited genetic studies
- It can be grown on a simple synthetic medium in the laboratory.
- It completes its life cycle in about two weeks.
- A single mating produces a large number of progeny flies.
- The sexes are clearly different: male and female flies are easily distinguishable.
- It has many types of hereditary variations that can be seen with low power microscopes.
5. Linkage and Recombination
- Morgan carried out several dihybrid crosses in Drosophila to study genes that were sex-linked; they were similar to Mendel's dihybrid crosses in peas.
- Example: he crossed yellow-bodied, white-eyed females with brown-bodied, red-eyed males, and intercrossed their F1 progeny.
- ★ Exam imp The two genes did not segregate independently of each other: the F2 ratio deviated very significantly from 9 : 3 : 3 : 1, the ratio expected for independent genes.
- Morgan's group knew these genes were on the X chromosome.
- When the two genes of a dihybrid cross were on the same chromosome, the proportion of parental gene combinations was much higher than that of the non-parental type.
- Morgan attributed this to the physical association, or linkage, of the two genes.
- Morgan's group also found that genes grouped on the same chromosome can be linked to different degrees.
- Some genes were very tightly linked and showed very low recombination (Cross A below).
- Others were loosely linked and showed higher recombination (Cross B below).
- ★ Exam imp The genes white and yellow were very tightly linked and showed only 1.3 per cent recombination.
- ★ Exam imp The genes white and miniature wing showed 37.2 per cent recombination.
- In the figure, dominant wild-type alleles carry a + sign as a superscript (y+, w+, m+).
| Feature | Cross A (y and w) | Cross B (w and m) |
|---|---|---|
| Genes | Yellow body, white eye | White eye, miniature wing |
| Parental types | 98.7% | 62.8% |
| Recombinant types | 1.3% | 37.2% |
| Strength of linkage | Very tight (genes close together) | Loose (genes far apart) |
Who coined the terms linkage and recombination?
How much recombination did white and miniature wing show?
Which F2 ratio did Morgan's linked genes fail to give?
6. Gene Maps
- ★ Exam imp Morgan's student Alfred Sturtevant used the frequency of recombination between gene pairs on the same chromosome as a measure of the distance between the genes.
- He used it to map the positions of genes on the chromosome.
- Today, genetic maps are widely used as a starting point in sequencing whole genomes, as in the Human Genome Sequencing Project.
- Closely located genes assort together; distantly located genes, because of recombination, assort almost independently.
- Linkage maps therefore correspond to the arrangement of genes on a chromosome.
- So the Law of Independent Assortment does not hold true for genes located on the same chromosome (linked genes).
7. Exam Essentials
Pairs to Match
| Scientist or term | Matches with |
|---|---|
| de Vries, Correns, von Tschermak | Rediscovered Mendel's results in 1900 |
| Walter Sutton | Chromosomal theory of inheritance |
| Sutton and Boveri | Parallel behaviour of chromosomes and genes |
| Thomas Hunt Morgan | Linkage and recombination in Drosophila |
| Alfred Sturtevant | Gene mapping by recombination frequency |
| Linkage | Physical association of genes on a chromosome |
| Recombination | Generation of non-parental gene combinations |
| White and yellow | 1.3% recombination; tight linkage |
| White and miniature wing | 37.2% recombination; loose linkage |
| Chromosomes | Coloured bodies seen by staining |
| Mitosis | Equational division |
| Meiosis | Reduction division |
| Genetic maps | Starting point for whole-genome sequencing |
- Linked genes do not follow the Law of Independent Assortment.
- Linked genes give a F2 that is not 9 : 3 : 3 : 1; parental types are in excess.
- Morgan's crosses studied sex-linked genes on the X chromosome, unlike Mendel's genes in peas.
- Sutton and Boveri gave the idea, but only Sutton named the chromosomal theory; Morgan verified it experimentally.
- Low recombination does not mean loose linkage; it means tight linkage.
Numbers to Remember
- 1865: Mendel published his work; 1900: rediscovery.
- 1902: chromosome movement during meiosis worked out.
- Drosophila life cycle: about two weeks.
- White-yellow: 1.3% recombination (98.7% parental).
- White-miniature wing: 37.2% recombination (62.8% parental).
8. Quick Revision
- Mendel published in 1865; ignored till 1900 for four reasons: communication, continuous variation, mathematics, no physical proof.
- 1900: de Vries, Correns and von Tschermak rediscovered his results independently.
- Chromosomes: double and divide before cell division; coloured bodies seen by staining.
- 1902: chromosome movement in meiosis worked out.
- Sutton and Boveri: chromosomes behave like genes; alleles sit at homologous sites on homologous chromosomes.
- Both occur in pairs, segregate at gamete formation and assort independently.
- Chromosome pairs align independently at the metaphase plate in meiosis I.
- Sutton named the chromosomal theory of inheritance.
- Morgan verified it using Drosophila melanogaster.
- Drosophila: synthetic medium, two-week life cycle, many progeny, distinct sexes, visible variations.
- Yellow-bodied, white-eyed females × brown-bodied, red-eyed males: F2 far from 9 : 3 : 3 : 1.
- Linkage: physical association of genes; recombination: non-parental combinations.
- White-yellow 1.3% (tight); white-miniature 37.2% (loose).
- Sturtevant: recombination frequency measures gene distance; gene maps.
- Genetic maps guide whole-genome sequencing, as in the Human Genome Project.
9. Solved Examples
List I: A. Sutton, B. Morgan, C. Sturtevant, D. de Vries
List II: I. Mapped genes using recombination frequency, II. Rediscovered Mendel's results, III. Chromosomal theory of inheritance, IV. Coined the terms linkage and recombination
Choose the correct answer:
(A) A-III, B-IV, C-I, D-II
(B) A-IV, B-III, C-I, D-II
(C) A-III, B-I, C-IV, D-II
(D) A-II, B-IV, C-I, D-III
Answer: (A). Sutton named the chromosomal theory (III), Morgan coined linkage and recombination (IV), Sturtevant mapped genes (I), and de Vries was one of the three who rediscovered Mendel (II).
A. He worked with Drosophila melanogaster.
B. The genes he studied in these crosses were on the X chromosome.
C. White and yellow showed 37.2 per cent recombination.
D. Parental gene combinations were much more frequent than non-parental ones.
E. His dihybrid F2 gave a 9 : 3 : 3 : 1 ratio.
Choose the correct answer:
(A) A, B and D only
(B) A, C and D only
(C) B, C and E only
(D) A, B, D and E only
Answer: (A). C is wrong: white and yellow showed 1.3 per cent; 37.2 per cent was for white and miniature wing. E is wrong: the ratio deviated very significantly from 9 : 3 : 3 : 1.
A. Rediscovery of Mendel's results
B. Mendel publishes his work
C. Chromosome movement during meiosis worked out
D. Mendel begins his hybridisation experiments
Choose the correct answer:
(A) D, B, A, C
(B) B, D, A, C
(C) D, B, C, A
(D) D, A, B, C
Answer: (A). Experiments began in 1856 (D), publication in 1865 (B), rediscovery in 1900 (A), and chromosome movement in meiosis by 1902 (C).
(A) The first pair is loosely linked
(B) The first pair lies closer together
(C) The second pair assorts in a 9 : 3 : 3 : 1 ratio
(D) Both pairs are unlinked
Answer: (B). Recombination frequency measures distance, so 2 per cent means the genes are close and tightly linked; 30 per cent means they are farther apart and loosely linked.
(A) It completes its life cycle in about two weeks
(B) A single mating gives many progeny
(C) Its sexes look identical
(D) It shows many hereditary variations visible under low power microscopes
Answer: (C). The sexes of Drosophila are clearly different and easy to tell apart.
10. Practice Questions
- Match List I with List II.
List I: A. Linkage, B. Recombination, C. Chromosomes, D. Meiosis
List II: I. Reduction division, II. Coloured bodies, III. Non-parental gene combinations, IV. Physical association of genes on a chromosome
(A) A-IV, B-III, C-II, D-I (B) A-III, B-IV, C-II, D-I (C) A-IV, B-III, C-I, D-II (D) A-II, B-III, C-IV, D-IAnswer: (A). Linkage-IV, recombination-III, chromosomes-II, meiosis-I. - Read the statements.
A. Chromosomes and genes both occur in pairs.
B. Alleles of a gene pair lie on non-homologous chromosomes.
C. Chromosome pairs align independently at the metaphase plate in meiosis I.
D. Mitosis is a reduction division.
Choose the correct answer: (A) A and C only (B) A, B and C only (C) B and D only (D) A, C and D onlyAnswer: (A). B is wrong (homologous sites on homologous chromosomes); D is wrong (mitosis is equational). - Which of the following was NOT a reason for Mendel's work remaining unrecognised? (A) Poor communication (B) Use of mathematics in biology (C) Lack of physical proof of factors (D) His experiments used too few plantsAnswer: (D). His experiments had a large sampling size.
- Statement I: Tightly linked genes show high recombination.
Statement II: The genes white and yellow in Drosophila are tightly linked.
(A) Both correct (B) I correct, II incorrect (C) I incorrect, II correct (D) Both incorrectAnswer: (C). Tightly linked genes show low recombination; white and yellow showed only 1.3 per cent. - In Cross B of Morgan (white and miniature wing), what percentage of the gametes of the F1 female carried parental combinations?Answer: 62.8 per cent (100 - 37.2).
- Two heterozygous parents are crossed. If the two loci are linked, what would be the distribution of phenotypic features in the F1 generation for a dihybrid cross?Answer: With linked loci, independent assortment fails. Parental combinations appear in much higher proportion than recombinant ones, so the progeny depart from 9 : 3 : 3 : 1. With complete linkage, the result depends on how the alleles sit: if both dominant alleles are on the same chromosome (AB/ab × AB/ab), the progeny are 3 dominant : 1 recessive, as for a single gene; if they are on opposite homologues (Ab/aB), they are 1 : 2 : 1. With loose linkage, some recombinants appear, in proportion to the distance between the genes.
- Briefly mention the contribution of T.H. Morgan in genetics.Answer: He experimentally verified the chromosomal theory of inheritance using Drosophila. His dihybrid crosses on X-linked genes showed that genes on the same chromosome do not assort independently. He coined the terms linkage and recombination, and showed that genes differ in the strength of their linkage (white-yellow 1.3%, white-miniature 37.2%).
- Who proposed the chromosomal theory of inheritance?Answer: Walter Sutton, who, with Theodore Boveri, noted the parallel behaviour of chromosomes and genes and united chromosomal segregation with Mendelian principles.
Common Mistakes to Avoid
- Saying Morgan proposed the chromosomal theory. Correct: Sutton proposed it; Morgan verified it experimentally.
- Writing 37.2% for white and yellow. Correct: white-yellow is 1.3%; white-miniature wing is 37.2%.
- Linking high recombination with tight linkage. Correct: tight linkage gives low recombination.
- Saying Mendel's work was ignored because of small samples. Correct: his samples were large; the reasons were communication, continuous variation, mathematics and no physical proof.
- Placing alleles on non-homologous chromosomes. Correct: alleles lie at homologous sites on homologous chromosomes.
- Expecting 9 : 3 : 3 : 1 from linked genes. Correct: linked genes give an excess of parental combinations.
- Calling mitosis a reduction division. Correct: mitosis is equational; meiosis is the reduction division.
- Crediting Morgan with gene mapping. Correct: his student Alfred Sturtevant mapped genes.
Frequently Asked Questions
Why did Mendel's work remain unrecognised for 35 years?
Communication was poor, so the work was not widely known. His idea of discrete, non-blending factors did not seem to explain the continuous variation seen in nature. Using mathematics in biology was unacceptable to many biologists, and he could not give physical proof of what factors were.
What is the chromosomal theory of inheritance?
It states that genes are carried on chromosomes, and that the pairing, segregation and independent assortment of chromosomes during meiosis explain Mendel's laws. Sutton and Boveri noted the parallel behaviour of chromosomes and genes, and Sutton named the theory. Morgan later verified it experimentally.
How does chromosome behaviour in meiosis explain Mendel's laws?
Homologous chromosomes, which carry the two alleles of a gene, separate in meiosis I, so each gamete gets one allele: this is segregation. Different chromosome pairs align at the metaphase plate independently, so genes on different chromosomes assort independently.
Why was Drosophila melanogaster suitable for Morgan's experiments?
It grows on a simple synthetic medium, completes its life cycle in about two weeks, and a single mating gives many progeny. Males and females are easy to tell apart, and it shows many hereditary variations that can be seen with low power microscopes.
What is the difference between linkage and recombination?
Linkage is the physical association of genes on the same chromosome, which keeps parental combinations together. Recombination is the generation of non-parental gene combinations. Morgan coined both terms. Tightly linked genes show little recombination, while loosely linked genes show more.
What did Morgan find for the genes white, yellow and miniature wing?
White and yellow were very tightly linked, with only 1.3 per cent recombination, so 98.7 per cent of combinations stayed parental. White and miniature wing were loosely linked, with 37.2 per cent recombination. So the linkage between yellow and white is stronger than between white and miniature.
How did Sturtevant make a genetic map?
Alfred Sturtevant, Morgan's student, used the frequency of recombination between pairs of genes on the same chromosome as a measure of the distance between them. From these distances he mapped the order and positions of genes on the chromosome.
Why do linked genes not follow the Law of Independent Assortment?
Genes on the same chromosome are physically associated, so they tend to pass together into the same gamete. Parental combinations therefore greatly outnumber recombinant ones, and a dihybrid cross does not give the 9 : 3 : 3 : 1 ratio expected for independent genes.
Previous year questions on Linkage and Crossing over
3 questions from past papers, each with a step-by-step solution.
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