Experiments in Plant Hybridization (1865)

By Gregor Mendel

              

              

 [1] Introductory Remarks

 

In this paper, I will describe experiments in which I artificially fertilized plants. When I got the same results over and over again, I did further experiments.  Some people spend their whole lives studying this branch of science.

To do this for your whole life requires lots of courage to keep going when it gets tough or boring.  To study that long is the only way to discover the secrets of evolution. 

This paper will now present the results of an experiment.  This experiment was done on a certain group of plants, for eight years until I reached a conclusion.  The reader must decide whether the results are correct or not.

 

 [2] Selection of the Experimental Plants

 

To have a successful experiment, you need to choose the right plants most suitable for the experiment.

The selection of the plants must be done very carefully so as to not risk anything or get incorrect results.  The plants must:

 

1. Have different traits.

2. The hybrids of the plants must, during the flowering period, be protected from bugs that could pollinate them and create incorrect results.

3. The hybrids and their offspring should always be able to produce seeds that can be grown into more plants.

           

If bugs did pollinate the plants, it would have a bad effect on the results.  If the plants could no longer produce seeds that could be grown into more plants, the experiments would be very difficult and it could even end them.  To determine the relationship between plants, the plants must be closely watched at all times.

34 different pea plants were gotten and were put into a two-year trial.  They were allowed to fertilize themselves and 22 were selected that always produced offspring that looked exactly like the parent plant.

 

[3] Division and Arrangement of the Experiments

 

Suppose you have two plants that are different for one particular characteristic. The goal of this experiment was to find out why, when the two plants are crossed, only one characteristic shows up.

 

The characteristics that were selected for these experiments were:

 

            1. The shape of the ripe seeds: either round or wrinkled.

2. The color of the inside of the seed: either pale yellow, bright yellow or orange colored.

            3. The color of the seed coat: either white or grayish brown.

            4. The shape of the ripe pods: either puffed up or wrinkled and squished.

5. The color of the unripe pods: either green or yellow. *

6. The position of the flowers on the stem: either distributed along the main stem or bunched at the top of the stem.

            7. The length of the stem: either tall or short.

 

A plant with each characteristic was crossed with a plant with the opposite characteristic, and the following numbers of fertilizations were done:

 

     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.

 

* One pea plant group had a brownish-red colored pod, which turned violet or blue when it ripened.

 

The plants were grown in garden beds, and a few in pots, and were kept upright using sticks, branches of trees, and strings stretched between sticks. For each experiment a number of potted plants were put in a greenhouse, to make sure the results were correct and to make sure that bugs didn’t pollinate them.

 

[4] The Forms of the Hybrids

 

In other experiments, it was found that if you crossed, for example, a tall plant and a short plant, the hybrids produced were not medium-sized, they were tall!  Peas are exactly the same.  Only one characteristic is shown.  The “shown” characteristic is called dominating and the “unshown” characteristic is called recessive. 

 

The characteristics that were dominant in these experiments were:

 

   1. The round shape of the seed.

   2. The yellow color of the inside of the seed.

   3. The grayish-brown color of the seed coat.

   4. The puffed up shape of the pod.

   5. The green color of the unripe pod.

   6. The distribution of the flowers along the main stem.

   7. The tall stem.

 

[5] The First Generation from the Hybrids

 

In this experiment, the hybrids produced in earlier experiments were allowed to self-fertilize.  In this generation the dominant traits reappeared, along with the recessive ones.  If there were four offspring plants, three would show a dominant characteristic and one would show a recessive characteristic, making a 3:1 ratio.

 

The characteristics that were recessive were:

1.     The wrinkled seed.

2.     The green color of the inside of the seed.

3.     The white color of the seed coat.

4.     The wrinkled, squished shape of the pod.

5.     The yellow color of the unripe pod.

6.     The flowers bunched at the top of the stem.

7.     The short stem.

 

    * Exp. 1:  Shape of the seed: 253 plants and 7324 seeds were gotten.  There were 5474 round seeds and 1850 wrinkled seeds.  The ratio was 2.96:1.

    * Exp. 2:  Color of the inside of the seed: 258 plants and 8023 seeds = 6022 yellow and 2001 green.  The ratio was 3.01:1.

 

              Experiment 1             Experiment 2

              Shape of Seed           Color of Inside of Seed

Plants Round Angular             Yellow Green

            1      45      12                25         11

            2      27       8               32           7

            3      24       7               14           5

            4      19      10              70           27

            5      32      11              24           13   

            7      88      24              32           13

            8      22      10               44           9

            9      28       6               50           14

           10     25       7               44           18

 

    * Exp. 3: Color of the seed coat:  Among 929 plants, 705 had grayish-brown seed coats and 224 had white seed coats, giving a ratio of 3.15:1.

    * Exp. 4: Shape of the pods:  Of 1181 plants, 882 of them had puffed up pods and 299 had wrinkled ones. The resulting ratio was 2.95:1.

    * Exp. 5: Color of the unripe pods:  428 had green pods and 152 had yellow ones, giving a ratio of 2.82:1.

    * Exp. 6: Position of flowers on the stem: Among 858 plants, 651 had flowers distributed evenly on the stem and 207 had flowers only at the top, giving a ratio of 3.14:1.

    * Exp. 7: Length of stem:  Out of 1064 plants, 787 had a tall stem, and 277 had a short stem, giving a ratio of 2.84:1.

 

If the results of the whole experiment were brought together, the average ratio was 2.98:1, or 3:1.

 

[6] The Second Generation from the Hybrids

 

            The plants from the previous experiment (the offspring produced when the hybrids self-fertilized) were allowed to self-fertilize.  The plants that showed the recessive characteristic produced offspring that were also recessive. Of those plants that had the dominant character, two thirds of them actually were hybrids and had both the dominant and recessive character, and one third had only the dominant character, for a ratio of 2:1.

 

Those plants that had only the dominant character produced offspring that all showed the dominant characteristic.  The remaining hybrid plants produced some offspring that showed the dominant characteristic and some that showed the recessive characteristic, in a ratio of 3 dominant to one recessive, just like when their hybrid parents were allowed to self-fertilize.

 

The separate experiments in which the offspring that showed the dominant characteristic were allowed to self-fertilize had these results:

 

    *  Exp. 1:  Among 565 plants that were raised from round seeds, 193 produced round seeds only, and 372 produced some plants with round seeds and some plants with wrinkled seeds, in a proportion of 3:1.

    *  Exp. 2:  Of 519 plants that were raised from seeds for which the inside of the seed was yellow, 166 produced seeds that were yellow inside and 353 produced some seeds whose insides were yellow and some seeds whose insides were green, in a ratio of 3:1.

    * Exp. 3:  The offspring of 36 plants produced only gray-brown seed coats, while some of  the offspring of 64 plants produced grayish-brown seed coats and some produced white seed coats..

    * Exp. 4:  The offspring of 29 plants produced only puffed up pods while some of the offspring of 71 plants had puffed up pods and some had wrinkled pods.

    * Exp. 5:  The offspring of 40 plants produced only green pods, while some of the offspring of 60 plants produced green  pods and some produced yellow ones.

    * Exp. 6.:  The offspring of 33 plants had only flowers distributed on the stem, while some of the offspring of 67 had flowers distributed on the stem and some had flowers bunched on the top.

*  Exp. 7:  The offspring of 28 plants had only tall stems, while some of the offspring of 72 plants had tall stems and some had short stems.

 

[7] The Subsequent Generations from the Hybrids

 

Summary:  The offspring produced when hybrids self-fertilized can be classified as either hybrids (possessing both the dominant and the recessive character), dominant (possessing only the dominant character), or recessive (possessing only the recessive character).  The ratio was 2 hybrids to one dominant to one recessive (2:1:1 ratio).

 

If A refers to one of the two constant characters, for instance the dominant character, and a refers to the recessive characters, and Aa refers to the hybrid form in which both characters are present, the expression

                               

                A + 2Aa + a

 

shows the proportion of offspring of the self-fertilizing hybrids.

 

[8] The Offspring of Hybrids in which Several Differentiating Characters are Associated.

 

Summary:  Experiments were done in which plants were crossed that differed in two or three characters.  For example, crosses were done among plants that differed in all three of the following characters:  seed coat form (round or wrinkled), color of the inside of the seed (green or yellow) and seed coat color (grayish-brown or white).

 

[9] The Reproductive Cells of the Hybrids

 

            Summary:  Each parent plant had two factors that were able to pass on traits from parent to offspring, but that only one of those factors was present in the egg cell or the pollen cell.  So hybrid parent plants produced eggs or pollen, and equal numbers of the egg or pollen cells had the dominant character and the recessive character.

 

Separate experiments proved that this was true of both pollen and egg cells.  It is purely a matter of chance, which of the two sorts of pollen will become united with each separate egg cell.

 

So when crossing two hybrids (Aa X Aa), the hybrids are able to produce, besides the two parental forms, offspring that are like themselves. A/a and a/A both give the same union Aa, since it makes no difference in the result of fertilization to which of the two characters the pollen or egg cells belong.   So A/A + A/a + a/A + a/a = A + 2 Aa + a.

 

 [10] Experiments with Hybrids of Other Species of Plants

 

Summary:  Other experiments were used to determine whether the laws discovered for peas applied to the hybrids of other plants.  Several experiments were done with different species of beans.

 

With some crosses of beans, looking at traits that determine the form of the plants, the ratios of the numbers in which the different forms appeared in the separate generations were the same as with peas.  But in other crosses with other species of beans, in which flower color was examined, the results were confusing.

 

 [11] Concluding Remarks

 

Summary:  With peas it was shown by experiment that the hybrids form egg and pollen cells of different kinds, and that this holds the key to the variability of their offspring.  So in peas, it is beyond a doubt that when a new embryo is formed, a perfect union of the elements of both reproductive cells must take place. How could we otherwise explain that among the offspring of the hybrids, both original types reappear in equal numbers and with all their peculiarities?

 

But even the truth of the law formulated for peas must still be confirmed and some of the more important experiments must be repeated.

 

It is important to compare the observations made regarding peas with the results arrived at by the two authorities in this branch of knowledge, Köreuter and Gärtner, in their investigations. A description follows of experiments carried out by Kölreuter, Gärtner, and others, including experiments made with respect to the transformation of one species into another by artificial fertilization.

 

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