Punnett square

The Punnett square is a square diagram that is

used to predict the genotypes of a particular
cross or breeding experiment. It is named after
Reginald C. Punnett, who devised the approach in
1905.[3][4][5][6][7][8] The diagram is used by
biologists to determine the probability of an
offspring having a particular genotype. The
Punnett square is a tabular summary of possible
combinations of maternal alleles with paternal
alleles.[9] These tables can be used to examine
the genotypical outcome probabilities of the
offspring of a single trait (allele), or when
crossing multiple traits from the parents. The
Punnett square is a visual representation of
Mendelian inheritance. It is important to
understand the terms "heterozygous",
"homozygous", "double heterozygote" (or
homozygote), "dominant allele" and "recessive
allele" when using the Punnett square method.
For multiple traits, using the "forked-line
method" is typically much easier than the
Punnett square. Phenotypes may be predicted
with at least better-than-chance accuracy using
a Punnett square, but the phenotype that may
appear in the presence of a given genotype can in
some instances be influenced by many other
factors, as when polygenic inheritance and/or
epigenetics are at work.

A Punnett square showing a typical test

cross. (green pod color is dominant over
yellow for pea pods[1] in contrast to pea
seeds, where yellow cotyledon color is
dominant over green[2]).

Punnett squares for each combination of

parents' colour vision status giving
probabilities of their offsprings' status,
each cell having 25% probability in
theory.
Zygosity
Zygosity refers to the grade of similarity
between the alleles that determine one specific
trait in an organism. In its simplest form, a
pair of alleles can be either homozygous or
heterozygous. Homozygosity, with homo relating
to same while zygous pertains to a zygote, is
seen when a combination of either two dominant
or two recessive alleles code for the same trait.
Recessive are always lowercase letters. For
example, using 'A' as the representative
character for each allele, a homozygous dominant
pair's genotype would be depicted as 'AA', while
homozygous recessive is shown as 'aa'.
Heterozygosity, with hetero associated with
different, can only be 'Aa' (the capital letter is
always presented first by convention). The
phenotype of a homozygous dominant pair is 'A',
or dominant, while the opposite is true for
homozygous recessive. Heterozygous pairs always
have a dominant phenotype.[10] To a lesser
degree, hemizygosity[11] and nullizygosity[12]
can also be seen in gene pairs.

Monohybrid cross
"Mono-" means "one"; this cross indicates that
the examination of a single trait. This could
mean (for example) eye color. Each genetic locus
is always represented by two letters. So in the
case of eye color, say "B = Brown eyes" and "b =
green eyes". In this example, both parents have
the genotype Bb. For the example of eye color,
this would mean they both have brown eyes. They
can produce gametes that contain either the B or
the b allele. (It is conventional in genetics to
use capital letters to indicate dominant alleles
and lower-case letters to indicate recessive
alleles.) The probability of an individual
offspring's having the genotype BB is 25%, Bb
is 50%, and bb is 25%. The ratio of the
phenotypes is 3:1, typical for a monohybrid
cross. When assessing phenotype from this, "3"
of the offspring have "Brown" eyes and only one
offspring has "green" eyes. (3 are "B?" and 1 is
"bb")
 Paternal
B b
Maternal
B BB Bb

b Bb bb

The way in which the B and b alleles interact

with each other to affect the appearance of the
offspring depends on how the gene products
(proteins) interact (see Mendelian inheritance).
This can include lethal effects and epistasis
(where one allele masks another, regardless of
dominant or recessive status).

Dihybrid cross
More complicated crosses can be made by looking
at two or more genes. The Punnett square works,
however, only if the genes are independent of
each other, which means that having a particular
allele of gene "A" does not alter the probability
of possessing an allele of gene "B". This is
equivalent to stating that the genes are not
linked, so that the two genes do not tend to sort
together during meiosis.

The following example illustrates a dihybrid

cross between two double-heterozygote pea
plants. R represents the dominant allele for
shape (round), while r represents the recessive
allele (wrinkled). A represents the dominant
allele for color (yellow), while a represents the
recessive allele (green). If each plant has the
genotype RrAa, and since the alleles for shape
and color genes are independent, then they can
produce four types of gametes with all possible
combinations: RA, Ra, rA, and ra.

RA Ra rA ra

RA RRAA RRAa RrAA RrAa

Ra RRAa RRaa RrAa Rraa

rA RrAA RrAa rrAA rrAa

ra RrAa Rraa rrAa rraa

Since dominant traits mask recessive traits

(assuming no epistasis), there are nine
combinations that have the phenotype round
yellow, three that are round green, three that are
wrinkled yellow, and one that is wrinkled green.
The ratio 9:3:3:1 is the expected outcome when
crossing two double-heterozygous parents with
unlinked genes. Any other ratio indicates that
something else has occurred (such as lethal
alleles, epistasis, linked genes, etc.).
Forked-line method

The forked-line method (also known as the tree

method and the branching system) can also solve
dihybrid and multi-hybrid crosses. A problem is
converted to a series of monohybrid crosses, and
the results are combined in a tree. However, a
tree produces the same result as a Punnett
square in less time and with more clarity. The
example below assesses another double-
heterozygote cross using RrYy x RrYy. As stated
above, the phenotypic ratio is expected to be
9:3:3:1 if crossing unlinked genes from two
double-heterozygotes. The genotypic ratio was
obtained in the diagram below, this diagram
will have more branches than if only analyzing
for phenotypic ratio.

There are also Punnett squares for

epistasis. In these cases the genotype
epistatic over the other genes hinders their
expression in the phenotype.
See also
Mendelian inheritance
Karnaugh map, a similar diagram used for
Boolean algebra simplification

References
1. Mendel, Gregor Johann (1866) [1865].
Versuche über Pflanzen-Hybriden (http://www.
bshs.org.uk/bshs-translations/mendel/2016?p
age=14) . p. 14. Archived (https://web.archive.
org/web/20210329112319/http://www.
bshs.org.uk/bshs-translations/mendel/2016?p
age=14) from the original on 2021-03-29.
Retrieved 2020-06-01. {{cite
book}}: |work= ignored (help)
2. Mendel, Gregor Johann (1866) [1865].
Versuche über Pflanzen-Hybriden (http://www.
bshs.org.uk/bshs-translations/mendel/2016?p
age=47) . p. 47. Archived (https://web.archive.
org/web/20210329112808/http://www.
bshs.org.uk/bshs-translations/mendel/2016?p
age=47) from the original on 2021-03-29.
Retrieved 2020-06-01. {{cite
book}}: |work= ignored (help)
3. Punnett, Reginald Crundall (1907). Mendelism
(2 ed.). London, UK: Macmillan. (NB. The 1905
first edition of this book does not contain the
Punnett square. In 1911, the third edition
gives a more thorough explanation.)
4. Edwards, Anthony William Fairbank (March
2012). "Punnett's square" (https://www.resea
rchgate.net/publication/221824531) .
Studies in History and Philosophy of Science
Part C: Studies in History and Philosophy of
Biological and Biomedical Sciences. 43 (1):
219–224.
doi:10.1016/j.shpsc.2011.11.011 (http
s://doi.org/10.1016%2Fj.shpsc.2011.11.
011) . PMID 22326091 (https://pubmed.n
cbi.nlm.nih.gov/22326091) . Abstract: "The
origin and development of Punnett's Square for
the enumeration and display of genotypes arising
in a cross in Mendelian genetics is described.
Due to R. C. Punnett, the idea evolved through
the work of the 'Cambridge geneticists',
including Punnett's colleagues William
Bateson, E. R. Saunders and R. H. Lock, soon
after the rediscovery of Mendel's paper in
1900. These geneticists were thoroughly
familiar with Mendel's paper, which itself
contained a similar square diagram. A
previously-unpublished three-factor diagram by
Sir Francis Galton existing in the Bateson
correspondence in Cambridge University Library
is then described. Finally the connection between
Punnett's Square and Venn Diagrams is
emphasized, and it is pointed out that Punnett,
Lock and John Venn overlapped as Fellows of
Gonville and Caius College, Cambridge."
5. Edwards, Anthony William Fairbank (September
2012). "Reginald Crundall Punnett: First
Arthur Balfour Professor of Genetics,
Cambridge, 1912" (https://www.ncbi.nlm.nih.g
ov/pmc/articles/PMC3430543) .
Perspectives. Genetics. Gonville and Caius
College, Cambridge, UK: Genetics Society of
America. 192 (1): 3–13.
doi:10.1534/genetics.112.143552 (http
s://doi.org/10.1534%2Fgenetics.112.14
3552) . PMC 3430543 (https://www.ncbi.
nlm.nih.gov/pmc/articles/PMC3430543) .
PMID 22964834 (https://pubmed.ncbi.nlm.n
ih.gov/22964834) . pp. 5–6: "[…] Punnett's
square seems to have been a development of
1905, too late for the first edition of his
Mendelism (May 1905) but much in evidence in
Report III to the Evolution Committee of the
Royal Society [(Bateson et al. 1906b)
"received March 16, 1906"]. The earliest
mention is contained in a letter to Bateson from
Francis Galton dated October 1, 1905
(Edwards 2012). We have the testimony of
Bateson (1909, p. 57) that "For the
introduction of this system [the 'graphic
method'], which greatly simplifies difficult
cases, I am indebted to Mr. Punnett." […] The
first published diagrams appeared in 1906.
[…] when Punnett published the second edition
of his Mendelism, he used a slightly different
format ([…] Punnett 1907, p. 45) […] In the
third edition (Punnett 1911, p. 34) he
reverted to the arrangement […] with a
description of the construction of what he
called the "chessboard" method (although in
 truth it is more like a multiplication table).
[…]" (11 pages)
6. Wimsatt, William C. (2012-05-15), "The
analytic geometry of genetics: Part I: the
structure, function, and early evolution of
Punnett squares", Archive for History of Exact
Sciences, 66 (66): 359–396 [359],
doi:10.1007/s00407-012-0096-7 (htt
ps://doi.org/10.1007%2Fs00407-012-0
096-7) , S2CID 119557681 (https://ap
i.semanticscholar.org/CorpusID:11955768
1)
7. Edwards, Anthony William Fairbank (June
2016). "Punnett's square: a postscript" (http
s://www.sciencedirect.com/science/article/ab
s/pii/S1369848616000157) . Studies
in History and Philosophy of Science Part C:
Studies in History and Philosophy of Biological
and Biomedical Sciences. Elsevier Ltd. 57:
69–70.
doi:10.1016/j.shpsc.2016.01.001 (http
s://doi.org/10.1016%2Fj.shpsc.2016.01.
001) . Retrieved 2021-03-29. (2 pages)
8. Müller-Wille, Staffan; Parolini, Giuditta
(2020-12-09). "Punnett squares and hybrid
crosses: how Mendelians learned their trade by
the book". Learning by the Book: Manuals and
Handbooks in the History of Science (https://w
ww.cambridge.org/core/journals/bjhs-themes/a
rticle/punnett-squares-and-hybrid-crosses-how
-mendelians-learned-their-trade-by-the-book/
18A1CE37A6EE536CC1CE1D4FF6FF31
74) . BJHS Themes. Vol. 5. British Society
for the History of Science / Cambridge
University Press. pp. 149–165.
doi:10.1017/bjt.2020.12 (https://doi.or
g/10.1017%2Fbjt.2020.12) .
S2CID 229344415 (https://api.semanticsc
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(https://web.archive.org/web/2021032911
1650/https://www.cambridge.org/core/journa
ls/bjhs-themes/article/punnett-squares-and-h
ybrid-crosses-how-mendelians-learned-their-tr
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E1D4FF6FF3174) from the original on
2021-03-29. Retrieved 2021-03-29. "
[…] Nilsson-Ehle was experimenting with a
visual arrangement that would become very
popular in Mendelian genetics. The lower half
of his notes comes close to what is known as the
'Punnett square' […] Punnett introduced this
square diagram to the literature in 1906 in a
paper co-authored with Bateson and Edith R.
Saunders, and included it in the second edition
of his Mendelism. In the third edition (1911),
he added a verbal description of how to
construct the diagram, and the Punnett square
became a standard feature of Mendelian
literature. As a detailed reconstruction by
 A.W.F. Edwards has shown, the diagram first
took shape in an exchange of letters between
Bateson and Galton for the more complex case of
a trihybrid cross, and may well have been
inspired by the way in which Mendel presented a
case of trifactorial inheritance of flower colour
in beans. […]"
9. Griffiths, Anthony J. F.; Miller, Jeffrey H.;
Suzuki, David T.; Lewontin, Richard C.; Gelbart,
William M. (2000). An Introduction to
Genetic Analysis (https://www.ncbi.nlm.nih.go
v/books/NBK22098/figure/A220/) (7 ed.).
New York, USA: W. H. Freeman.
10. AthenaMyth (2014-06-16).
"Dominant/Recessive vs Hetero/Homozygous" (h
ttps://athenamyth.deviantart.com/journal/Dom
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35429) . DeviantArt. Archived (https://web.a
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11. Shiel Jr., William C. (2018-12-12)
[2017]. "Medical Definition of Hemizygous"
(https://www.medicinenet.com/script/main/ar
t.asp?articlekey=3687) . MedicineNet.
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12. Robles, Ivan Suarez (2010-11-16).
"nullizygous" (https://web.stanford.edu/group/
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Further reading
Campbell, Neil Allison (2005). Biology
(7 ed.). Benjamin-Cummings Publishing
Company. ISBN 978-0-8053-7146-8.
OCLC 71890442 (https://www.worldcat.o
rg/oclc/71890442) .
External links
Online Punnett Square Calculator (https://dn
a.frieger.com/punnett.php)
Online Punnett Square Calculator, monohybrid
and dihybrid, autosomal and sex-linked (htt
p://www.bugaco.com/calculators/punnett_sq
uare.php)

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