The stamens are short; the grains of pollen smaller and oblong in shape. The upper half of the tube of the corolla is more expanded. The number of seeds produced is smaller and the ovules larger. The plants tend to flower first.

The short-styled plants have a short pistil, half the length of the tube of the corolla, with a smooth depressed stigma standing beneath the anthers. The stamens are long; the grains of pollen are spherical and larger. The tube of the corolla is of uniform diameter except close to the upper end. The number of seeds produced is larger.

I have examined a large number of flowers; and though the shape of the stigma and the length of the pistil both vary, especially in the short-styled form, I have never met with any transitional states between the two forms in plants growing in a state of nature. There is never the slightest doubt under which form a plant ought to be classed. The two kinds of flowers are never found on the same individual plant. I marked many cowslips and primroses, and on the following year all retained the same character, as did some in my garden which flowered out of their proper season in the autumn. Mr. W. Wooler, of Darlington, however, informs us that he has seen early blossoms on the Polyanthus, which were not long-styled, but became so later in the season. (1/4. I have proved by numerous experiments, hereafter to be given, that the Polyanthus is a variety of Primula veris.) Possibly in this case the pistils may not have been fully developed during the early spring. An excellent proof of the permanence of the two forms may be seen in nursery-gardens, where choice varieties of the Polyanthus are propagated by division; and I found whole beds of several varieties, each consisting exclusively of the one or the other form. The two forms exist in the wild state in about equal numbers: I collected 522 umbels from plants growing in several stations, taking a single umbel from each plant; and 241 were long-styled, and 281 short-styled. No difference in tint or size could be perceived in the two great masses of flowers.

We shall presently see that most of the species of Primula exist under two analogous forms; and it may be asked what is the meaning of the above-described important differences in their structure? The question seems well worthy of careful investigation, and I will give my observations on the cowslip in detail. The first idea which naturally occurred to me was, that this species was tending towards a dioecious condition; that the long-styled plants, with their longer pistils, rougher stigmas, and smaller pollen-grains, were more feminine in nature, and would produce more seed;--that the short-styled plants, with their shorter pistils, longer stamens and larger pollen-grains, were more masculine in nature. Accordingly, in 1860, I marked a few cowslips of both forms growing in my garden, and others growing in an open field, and others in a shady wood, and gathered and weighed the seed. In all the lots the short-styled plants yielded, contrary to my expectation, most seed. Taking the lots together, the following is the result:--

TABLE 1.1.

Column 1: Plant. Column 2: Number of Plants. Column 3: Number of Umbels Produced. Column 4: Number of Capsules Produced. Column 5: Weight of Seed In Grains.

Short-styled cowslips : 9 : 33 : 199 : 83. Long-styled cowslips : 13 : 51 : 261 : 91.

If we compare the weight from an equal number of plants, and from an equal number of umbels, and from an equal number of capsules of the two forms, we get the following results:--

TABLE 1.2.

Column 1: Plant. Column 2: Number of Plants. Column 3: Weight of Seed in grains. ... Column 4: Number of Umbels. Column 5: Weight of Seed. ... Column 6: Number of Capsules. Column 7: Weight of Seed in grains.

Short-styled cowslips : 10 : 92 :: 100 : 251 :: 100 : 41. Long-styled cowslips : 10 : 70 :: 100 : 178 :: 100 : 34.

So that, by all these standards of comparison, the short-styled form is the more fertile; if we take the number of umbels (which is the fairest standard, for large and small plants are thus equalised), the short-styled plants produce more seed than the long-styled, in the proportion of nearly four to three.

Charles Darwin

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