paper november 9

David Elihu
7.021 Section E
Changes to Paper
Made general corrections to grammar and sentence structure
Got rid of redundancies
Correctly labeled figures
David Elihu
7.021, Section E(W9-11)
Paper by Gregor Mendel, laws of segregation and independent assortment
There are currently a few different prevailing theories of inheritance.
One theory,
fathered by Aristotle, predicts that one parent is the majority contributor to the offspring’s
inherited features. Around the 5th century, Hippocrates published a collection of works on the
pattern of inheritance. He wrote, “the offspring resembles its parent because the particles of the
semen come from every part of the body." The theory, known as pangenesis, said that all parts
of an organism produce “gemmules,” which come together in the semen and are responsible for
the formation of the next organism. Another other theory contends that the traits of the parents
become mixed and are lost in the mixed traits of the offspring. This theory is called blended
inheritance. About 15 years ago, Abbot Napp proposed many questions about inheritance that
were never answered, such as how traits are inherited and whether or not chance is a determining
factor in passing along characteristics.
To this day no one has successfully attempted to explain the laws of inheritance, perhaps
because to do so would require Herculean effort. Previous experiments have led to inconclusive
results due to flaws in the experimental design. The results have not helped differentiate traits
inherited by offspring of the same parents. Nor have they isolated generations from one another.
Most important, there is no statistical data on the relationships between generations. We thus
have proposed an alternative theory of inheritance after choosing plants that are of different order
and share little if any traits with each other.
Our study aims to discover the general laws concerning the formation and development
of hybrids, and form a predictable method of artificial fertilization in order to obtain new
variations in offspring. It is hoped that this study, which spanned eight years, will provide a
better picture of the relationship between parent and offspring and help elucidate the laws that
govern inheritance.
Previous studies have demonstrated that if two plants that share virtually all
characteristics but a few are crossed, they will produce hybrids that share the common
characteristics but vary in the differentiating characters. The object of this experiment was to
determine the law by which these differentiating traits appear in successive generations of
Plants were chosen that had various traits and fertility throughout subsequent generations,
and whose natural pollination could be prevented.
The genus Pisum was chosen as the
experimental subject through careful comparison to other types of plants. Pisum Sativum, the
garden pea, was used in most of the experiments. Other types of pea plants that were chosen but
were used less frequently were P. quadratum, P. saccharatum, and P. umbellatum. In total, 34
varieties of pea plant were obtained. Note that the species denotations in these cases have a high
probability of being immaterial since one can not always distinguish between separate species or
mere varieties of the same species.
Plants from the genus Pisum created a favorable situation because the reproductive
organs are closely packed inside the keel and the anthers burst directly inside the bug. Through
this process, the stigma is covered with pollen before the flower even opens, thus preventing
pollination from an insect. The peas were also chosen as the experimental model because their
short growth time would speed up the retrieval of our results.
During the experiment, there were seven different characteristics of the peas that were
selected for the experiments. They are listed in table 1a below:
1. form of the ripe seeds
Round (possibly partially), with occasional
shallow depressions on the surface. Otherwise
they are irregularly angular and considerably
2. color of the seed endosperm
Pale yellow, bright yellow and orange, or a less
intense green tint. Since coats are transparent
can examine color
3. seed-coat color
White (correlate with white flowers); gray,
gray-brown, leather-brown, with or without
violet spotting. If without violet spotting, the
color of the standards is violet, the wings are
purple, and the stem in the axils of the leaves
has a reddish tint. When boiled in water, the
gray seed-coats become dark brown.
4. form of the ripe pods
Their shape is either dome shaped (convex) or
depressed (concave).
5. color of the unripe pods
Light to dark green, or intense yellow. The
stalks, leaf-veins and calyx help determine the
6. flower position
Axial, distributed along the main stem, or
terminal, where they gather at the top of the
stem. In the terminal case the upper part of the
stem is wider.
7. stem length
Vary considerably in length. To be able to
differentiate crossing occurred among plants
with widely varying stem lengths (long X short
Table 1. Differentiating characteristics used for the pea fertilizations. The peas chosen were
selected to show differences in length and color of the stem; size and shape of the leaves; in the
appearance of the flowers and pods; in the length of the flower stalk; in the form and size of the
seeds and the color of their coats; and in the color of the endosperm.
Crosses were done among plants with one trait that was differentiating between them.
Characteristics that were not visible different were acceptable since we were not able to control
for them. When there was more than plant that had the possibility of being cross-fertilized, the
healthier plant with stronger trait expression was chosen. In addition, if a plant was to be used to
cross two different traits, it would be the pollen-plant in one cross and the seed-bearer in the
The cross-fertilizations were conducted as follows:
Experiment (similar to trait number)
Fertilization Data
1st experiment
60 fertilizations on 15 plants
2nd experiment
58 fertilizations on 10 plants
3rd experiment
35 fertilizations on 10 plants
4th experiment
40 fertilizations on 10 plants
5th experiment
23 fertilizations on 5 plants
6th experiment
34 fertilizations on 10 plants
7th experiment
37 fertilizations on 10 plants
Table 2. Description of seven types of cross-fertilizations. The cross-fertilization numbers (17) are analogous to the differentiating characteristics.
For example, the first experiment
examined inheritance laws regarding the form of the ripe seed.
The plants were grown in a natural environment. Garden beds were used, and the plants
were held upright using sticks, tree branches and string. Occasionally the plants were grown in
pots for the controls.1
After crossing plants which differed in one character, plants were crossed that differed in
2 and 3 characteristics. In the first cross, the parental plants differed in the form of seed and in
the color of the endosperm. In the second cross, which was between plants with three dissimilar
Mendel was not thorough at all describing the involvement of controls in his experiments. He describes that they
were used, but neglects to include characteristics that might be relevant.
traits, the form of the seed, the color of the endosperm, and the color of the seed-coats were
cross-fertilized. After these two experiments, further crosses, though smaller in number, of the
same types of plants were done in order to retrieve a hybrid plant.
To learn more about the inheritance of traits in organisms with dissimilar characteristics,
a plant with round form and a yellow endosperm was crossed in four different instances with a
form-wrinkled plant with a green endosperm. Further analysis of the inheritance of traits for
hybrids was done in several other experiments where variety in traits was available. Parents with
different types of traits were crossed in order to determine the rate at which different traits are
passed down to an offspring.
Controls were used to make sure insects in the natural outdoor environment did not affect
the offspring of the pollinations. The beetle Bruchus pisi can pose a threat to the experimental
purity as its females are known to lay the eggs in the flower, thus opening the keel. Another
possible impurity might be introduced by partial exposure of the fertilizing organs of a weak and
dying plant, causing fertilization when it was not wanted. To prevent these potential problems,
equivalent pot plants were grown in a greenhouse to serve as a control.
Artificial fertilization, a relatively intricate process, made it possible to control of the
reproductive conditions of the peas.
In the process, the bud of a pea was opened before
development was complete, and the keel was removed. Next each stamen was carefully removed
with forceps, making sure it wasn’t punctured. Cross-fertilization was induced by dusting the
stigma over on the seed-bearing plant using pollen from the host plant.
First Generation
From this point forward, those traits that remain unchanged in the offspring of the hybrid
cross will be known as dominant. Those that become hidden in the process of a cross will be
known as recessive. Throughout the experiments, it was shown that whether the dominant trait
is contained within the seed plant or the pollen plant does not affect the outcome of the hybrid
crosses. The dominant characteristics in the differentiating crosses were determined to be as
The round or roundish seed with or without shallow depressions.
The yellow coloring of the seed abdomen.
The gray, gray-brown, or leather brown color that lies in the seed coat, in
conjunction with red-violet blossoms and reddish spots in the leaf axils.
the inflated form of the pod
the green coloring of the unripe pod in conjunction with the stems, leaf-veins and
calyx all having the same color.
The distribution of flowers along the stem.
The longer length of the stem.
These results were all determinable within the first generation of crosses between the species.
In this first generation, the ratio of dominant to recessive recurrence of traits appeared in a 3:1
ratio. This was observable in virtually all of the first generation crosses without exception. The
recurrence of the 3:1 ratio can be witnessed in the table below:
Total number
Total number
Number of
Number of
Ratio of
of hybrids
of seeds
Dominant to
1. form of the 253
ripe seeds
2. color of the 258
seed-coat Unknown
4. form of the Unknown
ripe pods
5. color of the Unknown
unripe pods
flower Unknown
7. stem length
Table 3. Results of cross-fertilizations. The predicted 3:1 ratio of dominant to recessive is
demonstrated in the results obtained. When available, the total number of hybrids is reported for
each cross. In addition, the total number of seeds, number of dominant, and number of recessive
pea plants is reported. From the dominant and recessive data, the ratios are determined.
Note that these results were obtained over a wide population survey. In more specific
cases, there were drastic fluctuations in individual small crosses, but once these data were
averaged out the above results were obtained. It should also be noted that in experiment 7,
where experiments were done for the length of the stem, the shorter plants that were grown were
immediately transferred to areas where they were not close to taller-stemmed plants, since the
taller plants would just overgrow them in neighboring areas.
Second Generation
When the plants with the recessive forms of the traits were crossed with each other, the
second generation results proved to be the same as the first generation results. The offspring
remained constantly recessive in the particular trait that was being followed. The same could not
be said for the dominant forms that were self-fertilized.
In the dominant crossing of experiment 1, of a total of 565 plants that came from the
round seeds of the first generation, 193 had round seeds only and were the same as the parental
generation. However, 372 plants gave both round and wrinkled seeds, in the proportion of 3:1.
The overall ratio was thus 1.93:1 for hybrids versus constants.
In experiment 2, the test of the color of the seed endosperm, a total of 519 plants were
grown. 166 yielded only the dominant yellow albumen, while 353 yielded both yellow and
green in the proportion of 3:1. The overall ratio for experiment 2 was 2.13:1.
Experiments 3-7 all had a population of 100 plants.
In experiment 3, 36 plants had only gray-brown seed coats, while 64 had some graybrown and some white. The ratio for experiment 3 was 1.78:1.
In experiment 4, 29 plants had only the dominant simply inflated pods, while 71 had a
mix of inflated and constricted. The ratio for experiment 4 was 2.45:1.
In experiment 5, 40 plants had green pods while 60 had both green and yellow pods. The
ratio for experiment 5 was 1.5:1. On a second trial, 35 had green pods while 65 had both green
and yellow pods. This ratio was 1.86:1.
In experiment 6, 33 plants had only axial flowers while 67 had some axial and some
terminal flowers. The ratio for experiment 6 was 2.03:1.
In experiment 7, 28 plants had long stems, while 72 had a mix of long and short stems.
The ratio for experiment 7 was 2.57:1.
This let us conclude that of the plants that showed the dominant trait in the first
generation, two-thirds were of the hybrid character, yielding a mix of traits in the second
generation; the other one-third kept the dominant trait in successive generations. Going back to
the original ratio of 3:1 in dominant to recessive show of traits, this ratio can be further refined
into a 2:1:1, which includes 2 hybrid dominant, 1 dominant, and 1 recessive in their traits.
Subsequent Generations
When further crosses were carried through, the results from above were confirmed.
Experiments 1 and 2 were carried through 6 generations, experiments 3 and 7 were carried
through 5 generations, and experiments 4, 5, and 6 were carried through 4 generations. The
offspring of the hybrids consistently segregated into the ratio of 2:1:1 determined above.
The Offspring of Hybrids in Which Several Differentiating Characters are Associated
When parental plants having more than one differentiating characteristic were crossed,
the offspring tended to approach the appearance of the parental plants which had the greater
number of dominant characters. When plants that differed in the form of seed and in the color of
the albumen were crossed, the following results were obtained. 556 seeds total were conceived
by 15 plants. Of the 556, 315 were round and yellow, 101 were wrinkled and yellow, 108 were
round and green, and 32 were wrinkled and green.
The round and yellow seeds were cross-fertilized again, and 38 had round yellow seeds,
65 had round yellow and green seeds, 60 had round yellow and wrinkled yellow seeds, and 138
had round yellow and green, wrinkled yellow and green seeds.
Of the wrinkled yellow seeds 96 plants resulted, of which 28 had only wrinkled yellow
seeds and 68 had wrinkled yellow and green seeds. Of the round and green seeds 102 plants
fruited, where 35 had only round green seeds and 67 had round and wrinkled green seeds. The
wrinkled green seeds fruited 30 plants which bore seeds all of the same character as their parents.
From these results, it was seen that 33 times there was a group seen that has constant
traits over time and does not vary in continuing generations. 65 times it was seen that there was
a group that had one hybrid with one dominant trait that continued throughout the generations.
138 times the plants were hybrid in both characters, and behaved just like the parents in their
own offspring. This resulted in an overall ratio of 1:2:4.
When similar experiments were carried out using a larger number of differentiating
characteristics, similar results were obtained. It was seen that the offspring of the hybrids which
have in between them several differentiating characteristics exhibit the terms of a series of
combinations, in which the offspring for each pair of differentiating traits are united.
Through these experiments, it was seen that the crossing between these plants was
essentially a crossing between a pollen and egg cell.