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Part 1: Understanding genotype and phenotype

By Maurice A. Dow, Ph.D.

Region 4, Ontario, Canada
One hundred and ten years ago Gregor Mendel's
work, originallypublished in 1866, was rediscovered
by three botanists and plant genetics blossomed. In
Here is an example of the range of phenotypes possible from one cross. (From left)
the time since, our understanding has advanced
'Elderberry Candy' (Stamile, 2003) and tetra 'Peppermint Delight' (Carpenter..].,
enormously, but, as always, there is still much to be
learned. Fifty years ago daylilies were being studied 2003) are the parents of 'Giant Panda' (Stamile, 2008) and 'Ruby Storm' (Stamile,
by scientific researchers. Unfortunately, genetics has 2008). The siblings are both early blooming six inch cultivars, but 'Giant Panda' is
since concentrated on the study of a few "model" an evergreen while 'Ruby Storm' is a dormant.
species and little modem research is done on - Photos courtesy of Pat Stamile except where noted
daylilies.
In 1972 Joanne orton--' hoped that in the future the complicat .. tinct and easily separable from each other) different and little aff ct..
ed daylilyflower color genetics would be understood well enough for ed by the environment. Those characteristics make the pea a go d
a geneticist to be able to look at a flower and indicate the gene ubject for genetic studies.
determining its color. However, daylilies are primarily self..incompatible (infertile wh n
Unfortunately, this is not possible yet for any plant species, even self..pollinated) perennials derived from crosses of many diff rent
those for which flower color has been studied intensively. Research species. Unfortunately, those are characteristics which make diploid
on many plant species has determined that flower color cannot con .. daylilies a difficult subject for genetic studies.
sistently predict the pigments present or the underlying genotypes. The foun ation for the study of genetics is:
Two flowers may have the same color but different pigments and Phenotype = Genotype + Environment + Genotyp X
therefore genotypes, or two flowers with the same pigment may have Environment Interaction + Error.
different colors".
Some work has been done with daylilies and much of the work Figure 1 ( ee next page) graphically show the effects of g n typ ,
done with other species such as petunias can be applied to daylilie . nvironment and their interaction in determining phen type (char..
In this series of articles, I will review the information which i acteri tic). If we ob erve everal cultivar (g notype ) in a ingle
available for daylilies. Research on the genetics of other pecies environment (temperature), we can see only the effect of gen type.
which is relevant to understanding the inheritance of variou trait If we ob erve a ingle cultivar in everal environment w can
in daylilies also will be included. only the effect of environment. If we observe several cultivars in v..
Daylily genetics is complicated, and there are several rea n for eral environment w can then ee the effect f gen type, envir n..
this. Mendel carefully chose the plant that he worke with - th ment and their interaction.
common garden pea. He spent several years making certain that th Cultivars E and F have a phenotype that i determine only by
trains he had were "pure ..breeding" (homozygous) for the trait h genotype an environm nt - there i no interacti n pre ent inc
was investigating. All the strains belonged to one annual dipl id their line are parallel. Cultivars A and G have a phen typ that i
plant pecies. Peas are self..compatible and naturally elf..fertilizing, almo t completely determined by genotype alon - there i no
and so each strain was completely inbred. The traits that he cho e t interaction and the effect of environment i negligible. Th remain..
rudy were well..defined, qualitatively (sharply di continuous, di .. ing cultivar B, C, 0, have phenotype that are determin by the
DJ .p.rin 1Q

•
.....-....
Q)
I"-
c..
~
+oJ
o
C
Glossary
Q)
..c Additive - all phenotypes have an additiv compo..
o, G
'-'"
nent a it represents the average of the two parental val . .
c ue. When the F 1 offspring, of a cro between a
o
:,t:j h mozygou plant howing a characteristic and a
CO
L-
F h mozyg u plant not h wing the characteri tic, h w
~
C E the characteri tic the phen type may be additive r
(],)
U D dominant r partially both.
C
o C Allele - a variant of a g ne. Alleles have ne or m r
o difference in their D A equence. atural genetic
....,
C B variati n in plant population i present a multiple alle . .
Q)
E
0>
A---- - - - - - - ---- - le for mo t genes. An allele of a particular gen may
have a very large effect on the phenotype, cau ing com..
a: pl te los of the function f the gene or it may have a
maller effect or no effect on the phenotype. Few allel
will have larg effects on a phenotype hile most will
Low Medium High have a mall or no mea urable effect. In a diploid indi..
Temperature (Environment) vidual each gene ha two all les which may be the am
( ee h mozygou ) or different (see heterozygou ).
Figure 1: The effects of genotype, environment and their Dominant - the dominance component of a phen . .
interaction in determining phenotype (characteristic) type is the difference between the ave rag of the tw
The hypothetical effect of genotype, environment and genotype X environ.. homozyg u parental ph notype and that of the F 1 off..
ment interaction on the phenotype using flower pigment concentration as an pring. When the dominance component i exactly
example phenotype and temperature as an example environmental factor. All equal to the additive component the phenotype i com..
hypothetical examples are assumed to follow accepted scientific practices in pl tely dominant. When the dominance component i
theirdesign and analysis. Seven different cultivars are represented by (A, B, C, 1 than the additive component partial d minance i
D, E, F). The symbols represent the averages of a number of measurements of pre ent r the dominance i incomplete. When th
pigment concentrations for each cultivar in each environment. The vertical d minance comp nent i zero the phenotype i perfect..
lines with arrows shown on the value for pigment concentration for culti ar G ly additive. Few phenotype will be perfectly additiv or
at low temperature represents the range of pigment concentrations shown by c mplet ly dominant. At the molecular le el of g n
the cultivar at that temperature. The ranges have been omitted from all other xpr ion u ually both allele in a diploid will
averages for clarity. ex re ed.
Genotype - the equence of the D A making up th
g n of an indi id ual.
c mbination f genotype, environ.. Heterozygous - in a dipl id the pre enc of two dif..
m nt and th ir int raction . The ferent allel for one gene. E.g. W /w individual are het..
lin s are not parallel and they even rozygou . The term for a tetraploid are different.
cr . . over. Becau e of the presenc Homozygous - in a diploid the pre enc of two iden..
of int raction the rder of th culti.. tical allele for ne gene. E.g. both W!W and w/w indi..
var i not th arne in the three vidual are h mozygou .
environment. We cann t gues Mutation - a change in the equence f th D A of
which cultivar will hav the mo t or a g ne. Thi r ult in a diff rent allele. It mayor may
th I ast pigment by I oking at them n t r ult in a different phen type. Mutati n may be
in only one environment. We al 0 I ctiv Iy n utral and have n mea urable effect on a
cannot gue how the envir nment plant, or th y may be deleteriou and 10 t by natural
will affect any specific cultivar (six 1 ction or b advantag ou and increa e by natural
f even cultivars have less pigment election to r place di advantag ou all le . Some alle . .
in high temp ra tu re than in medi . . le may form what are called balanced polymorphism in
urn temperature but cultivar B has which ca two or m re all le have advantage and di . .
more pigmen ). advantag in different circum tances, n ne i c n i . .
In the garden r field we 0 s rv t ntly b tt rand th allel are maintain d in the pop..
r measur phenotyp s and attempt ulati n at int rmediate frequencie. Mutation with
to d t rmin (gue ) the gen typ vi i 1 ff ct ccur very rar ly, nee in a milli n or once
fr m tho e ob ervation . To d 0 in a hundr d th u and.
we try to mak the environm nt Phenotype - rh ob erved characteri tic of an indi..
c n tant and beau e of amp ling vi ual, for xampl , mea ured height or flower color.
(the plants analyzed are only a sam . . Reces ive - a phenotyp that is completely masked by
pl of the infinite number of plant an alternativ ph notype.
with tho e characteristics that could
See Geneti c , page 14
~14~
Red F1 0ffspring
(Paren~)
The effect of complete and incomplete (partial) dominance and
additive phenotypes on visible flower colour: red and white. The value
of the additive component is represented by d and the value of the
Genetics
continued from page 13
opposite of this in school.
As an imaginary example, a normal plant
has red petals. If we find a naturally occur ..
ring mutation (new phenotype that is inher..
ited) or create a mutation in the lab that has
white petals (not known in daylilies), then
the mutation is named white with the gene
symbol w if the white phenotype is recessive.
A w/w plant would then be white, and a
W/W plant would be red.
As shown in Figure 2, if the cross of a
W/W (red) plant with a w/w (white) plant
produces red offspring (w/w) that are indis..
tinguishable from their red parent, then red
is described as being completely dominant to
white.
If the red offspring are paler than the red
parent, there is incomplete dominance.
When the color is halfway between red
and white, it is an average of the parents'
contribution (this is referred to as additive
and indicates no dominance).
If the color of the seedling is more red
than halfway between red and white, but still
not as red as the parent, this is the result of
partial red dominance. If the color of the
seedling is less red than half..way between
red and white but not as white as the white
parent then this is the result of partial domi..
nance for white. In most cases phenotype
will be additive with partial dominance
(more than halfway towards one of the par..
ents) rather than completely dominant or
perfectly additive.
Mendel worked with the genetics of flower
color within one species and found effects
determined by a single gene. Geneticists
have since found that the genetics of a char..
acteristic such as flower color is often simpler
when studied within a single species and
becomes much more complex when studied
in the offspringof crosses between two differ..
ent species. While a characteristic within a
Figure 2
Phenotype
Pigment Concentration
.......r----------d--------~
Mid-point
Pale red
dominance component is represented by h. PI and P2 are the parents
and F1 represents the first generation offspring from a cross of the two
parents.
species may be due to differences in one
gene, when between different species it may
be due to many genes>.
Daylilies originate from crosses between
many species and we will find that most
characteristics in daylilies are not inherited
simply but are due to differences in many
genes. In working within a species Mendel
determined that particular crosses produced
offspring showing defined ratios of the phe..
notypes. Second generation crosses (Fz)
showed the ratio 3: 1 for the dominant, reces..
sive phenotypes, for example. Research has
found that occasionally within a species
these ratios are not found when expected for
single genes but that in plants derived from
interspecific (between two or more species)
crosses these ratios are often very different
from those expected by Mendelian genet..
ics6.When this occurs it is called segregation
distortion. We can expect that segregation
distortion will be present in many daylily
crosses, and it will make it very difficult to
analyze crosses genetically or to predict the
numbers and types of offspring from crosses.
It is also important to understand that
how we measure a characteristic has a pro..
found effect on how we describe the inheri ..
tance of that characteristic. This is particu ..
lady important when making observations
on color. As an example, if we examine a
number of different cultivars for flower color
and characterize them by unaided eye as
being either red or not ..red, we might then
find that the results of crosses suggest that
red is dominant and a single gene. However,
if we reclassify the red..flowered plants by
measuring the amount of pigment in the
flowers, the results of the crosses might sug..
gest that red is additive and that more than
one gene is involved.
Unaided human vision is not very accu..
rate at determining differences when pig..
ments are present in high concentrations7or
DJ
=
h
d
dominance value
=additive value
White
(Parent 1)
in determining their presence or absence
when they are at low concentrations. This
can affect the conclusions made about the
inheritance of color characteristics since
phenotypes may be misclassified. A quanti..
tative (continuous phenotypic variation)
measure will usually be more representative
of the actual gene action since at the molec..
ular level both alleles (alternative forms of a
gene) will be expressed for most genes.
Editor's note: The next installment will be:
Pathways to color: From genotype to phenotype
References
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genetics. The HemerocallisJournal. 26 (2):29..
39, 1972.
zNorton , ]. Some basic Hemerocallis
genetics - Part 2. The Hemerocallis Journal.
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3Norton , ]. Some basic Hemerocallis
genetics - Part 3. The Hemerocallis Journal
27 (1):20..27, 1973.
4Griesbach, R. ]. The inheritance of flower
color in Petunia hybrida Vilm. Journal of
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5Dole, J. and R. Kessell. Inheritance of
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Accumulation, Induces Male Sterility, and Is
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