text:
RECORD: Darwin, C. R. 1864. On the sexual relations of the three forms of Lythrum salicaria. [Read 16 June] Journal of the Linnean Society of London (Botany) 8: 169-196, 1 text figure.
REVISION HISTORY: Scanned, OCRed, corrected and edited by John van Wyhe 2003-8, textual corrections by Sue Asscher 1.2007. RN3
NOTE: See manuscript drafts or notes on this item in DAR27.2
On the Sexual Relations of the Three Forms of Lythrum
By CHARLES DARWIN, F.R.S., F.L.S., &c.
[Read 16 June 1864.]
SOME of the species of Lythrum offer in their manner of fertilization a more remarkable case than can, perhaps, be found in any other plant or animal. In Lythrum salicaria three plainly different forms occur: each of these is an hermaphrodite, each is distinct in its female organs from the other two forms, and each is furnished with two sets of stamens or males differing from each other in appearance and function. Altogether there are three females and three sets of males, all as distinct from each other as if they belonged to different species; and if smaller functional differences are considered, there are five distinct sets of males. Two of the three hermaphrodites must coexist, and the pollen be carried by insects reciprocally from one to the other, in order that either of the two should be fully fertile; but unless all three forms coexist, there will be waste of two sets of stamens, and the organization of the species, as a whole, will be imperfect. On the other hand, when all three hermaphrodites coexist, and the pollen is carried from one to the other, the scheme is perfect; there is no waste of pollen and no false co-adaptation. In short, nature has ordained a most complex marriage-arrangement, namely a triple union between three hermaphrodites,—each her-
1 Purple loosestrife. Edited portions of this article appeared in Forms of flowers pp. 137-67. Darwin also discussed Lythrum salicaria in Origin 4th edn p. 323 and Variation 2: pp. 166, 183. See Correspondence vol. 12.
maphrodite being in its female organ quite distinct from the other two hermaphrodites and partially distinct in its male organs, and each furnished with two sets of males.
The three forms may be conveniently called, from the unequal lengths of their pistils, the long-styled, mid-styled, and short-styled. Their existence and differences were first observed by Vaucher*, and subsequently more carefully by Wirtgen;1 but, not being guided by any theory, neither author perceived some of the most curious points of difference. I will first briefly describe the three forms by the aid of the accompanying accurate diagram, which shows the flowers, six times magnified, in their natural position, with their petals and the near side of the calyx removed.
Long-styled form.—This can at once be recognized by the length of the pistil, which is (including the ovarium) fully one-third longer than that of the mid-styled, and more than thrice as long as that of the short-styled form. It is so disproportionately long, compared with the flower, that it projects in the bud through the unfolded petals. It stands out considerably beyond the longer stamens; its terminal portion depends a little, but the stigma itself is slightly upturned: the globular stigma is considerably larger than that of the other two forms. The six longer stamens project about two-thirds of the length of the pistil, and correspond in length with the pistil of the mid-styled form. The correspondence with the pistil in length in this and the two following cases is generally very close; the difference, where there is any, being usually in a slight excess of length in the stamens. The six shorter stamens (each of which alternates with a longer one) lie concealed within the calyx; their ends are upturned, and they are graduated in length, so as to form a triple row—both which characters are here much more marked than with the longer stamens, which vary in these respects. The anthers of the shorter stamens are smaller than those of the longer stamens. Knowing that the pollen differs greatly in the longer and shorter stamens of the two other forms, I carefully compared that of the two sets of stamens in this form: in both the pollen-grains are yellow, but they are a little larger in the longer than in the shorter stamens. The difference is slight, so that I convinced myself of its reality only by putting two small heaps close together under the compound mi-
* Hist. Phys. des Plantes d'Europe, tom. ii., 1841, p. 371.2 Wirtgen, "Ueber Lythrum salicaria und dessen Formen," Verhand. des naturhist. Vereins der preuss. Rheinl., 5. Jahrgang, 1848, S. 7.3
1 Philipp Wilhelm Wirtgen (1806-1870), German teacher and botanist who lived in Koblenz.
2 Vaucher 1841.
3 Wirtgen 1848.
Diagrams of the flowers of the three forms of Lythrum salicaria, in their natural position, with the petals removed and with the near side of the calyx cut away: enlarged six times.
The dotted lines with the arrows show which pollen must be applied to each stigma to cause full fertility.
croscope, and I found I could always (with one exception) distinguish them: I then showed the specimens to two other persons, and they likewise distinguished the two kinds and pointed out which was the largest. The capsules of this form contain, on an average, 93 seeds: how this average was obtained will presently be explained. I repeatedly observed that the seed, when cleaned, seemed larger than that from the mid-styled or short-styled forms; consequently I placed 100 long-styled seeds in a good balance, and by the double method of weighing found that they equalled 121 seeds of the mid-styled and 142 of the short-styled; or, in short, that five long-styled seeds equalled six mid-styled and seven short-styled seeds. These slight differences in the weight of the seed, and, as we shall soon see, in the average number produced, are worth recording, as they characterize not mere varieties but coexisting forms of the same species.
Mid-styled form.—The pistil occupies the position represented in the diagram, with its extremity considerably, but in a variable degree, upturned; the stigma is seated between the anthers of the long and the short stamens. The six longer stamens correspond in length with the pistil of the long-styled form; their filaments are coloured bright pink; the anthers are dark-coloured, but from containing bright green pollen and from their early dehiscence they appear emerald-green. Hence the general appearance of these stamens is remarkably dissimilar from that of the longer stamens of the long-styled form. The six shorter stamens, enclosed within the calyx, resemble in all respects the shorter stamens of the long-styled form, and both correspond in length with the short pistil of the short-styled form. The green pollen-grains of the longer stamens are plainly larger than the yellow pollen-grains of the shorter anthers: this fact was conspicuous in several camera-lucida drawings made for me by my son, Mr. W. E. Darwin.1 There is some variability in size, but 12/7000 of an inch may be taken as about the average diameter of the green pollen-grains when distended with water, and 9/7000 as the diameter of the yellow grains of the shorter stamens; so that the difference in diameter is in about the proportion of four to three. The capsules contain, on an average, 132 seeds; but, perhaps, as we shall see, this is rather too high an average. The seeds themselves are smaller than those of the long-styled form.
Short-styled form.—The pistil is here very short, not one-third of the length of that of the long-styled form. It is enclosed within the calyx, which, differently from in the other two forms,
1 William Erasmus Darwin (1839-1914), Darwin's eldest child. The drawings survive in DAR27.2.
does not enclose any anthers. The end of the pistil is generally bent upwards at right angles. The six longer stamens, with their pink filaments and green pollen, resemble in size of the grains and in all respects the longer stamens of the mid-styled form, and both correspond in length with the long-styled pistil. The six shorter stamens, with their uncoloured filaments and yellow pollen, resemble in size of the grains and in all respects the longer stamens of the long-styled form, and both correspond in length with the mid-styled pistil. The capsules contain fewer seeds on an average than in either of the preceding forms, namely 83.5, and they are considerably smaller in size. In this latter respect, but not in number, there is a gradation parallel to that of the length of the pistil, the long-styled having the largest, the mid-styled the next in size, and the short-styled the smallest seed.
From this description we see that there are three distinct female organs, or rather females as they are borne on distinct individuals, differing in the length and curvature of the style, in the size of the stigma, and in the number and size of the seed. In the three forms, taken together, there are thirty-six stamens or males, and these can be divided into three sets of a dozen each, differing from each other in length, curvature, and colour of the filaments, in the size of the anthers, and especially in the colour and diameter of the pollen-grains. Each of the three forms bears half-a-dozen of one kind of stamens and half-a-dozen of another kind, but not all three kinds. The three kinds correspond in length with the three pistils: the correspondence is always between half the stamens borne by two forms with the pistil of a third form. These remarks apply to the structure, and not, as yet, to the functions, of the reproductive organs.
I ascertained the average number of seed by counting them in eight fine selected capsules taken from plants of the three forms growing wild, and the result was, as we have seen, for the long-styled (neglecting decimals) 93, mid-styled 132, and short-styled 83. I should not have trusted this result, but I had a number of plants in my garden which, from their youth, did not yield the full complement of seed, but they were of exactly the same age and grew under exactly the same conditions, and were freely visited by bees. I took six fine capsules from each, and found the average to be for the long-styled 80, for the mid-styled 97, and for the short-styled 61. Lastly, I made numerous artificial unions, and, as may be seen in the following Tables, these gave in the long-styled an average of 90 seeds, in the mid-styled 117,
and in the short-styled 71. So that we have good concurrent evidence of the different average production of seed by the three forms. To show that the artificial fertilizations, presently to be described, produced their full effect and may be trusted, I may state that one mid-styled capsule yielded 151 good seeds, which is the exact number of the finest wild capsule examined by me. Artificially fertilized short- and long-styled capsules actually produced a greater number of seeds than I have found in wild plants, but then I did not examine many of the latter. This Lythrum, I may add, offers a remarkable instance, how profoundly ignorant we are of the life-conditions of each species: naturally it grows "in wet ditches, watery places, and especially on the banks of streams," and though it produces so many minute seeds, it never spreads on the adjoining land; yet, planted in my garden, on clayey soil lying over the chalk, and which is so dry that a rush cannot be found, it thrives luxuriantly, grows to above six feet in height, produces self-sown seedlings, and (which is a severer test) is fully as fertile as in a state of nature. Nevertheless it would be almost a miracle to find this plant spontaneously growing on such land as my garden, though under its native climate.
According to Vaucher and Wirtgen, the three forms coexist in all parts of Europe. Some friends gathered for me in North Wales a number of twigs from separate plants growing near each other, and then classified them. My son did the same in Hampshire, and here is the result:—
|North Wales
|Hampshire .
|Total . . . .
If twice or thrice the number had been collected, probably the three forms would have been found nearly equal; I infer this from considering the above figures, and from my son telling me that if he had collected in another spot, he felt sure that the mid-styled plants would have been in excess. I several times sowed small parcels of seed, and raised all three forms; but I neglected to record the parent form, except in one instance, in which I raised from short-styled seed twelve plants, of which only one turned out long-styled, four mid-styled, and seven short-styled.
Insects are necessary for the fertilization of this Lythrum. During two years I kept two plants of each form protected, and in the autumn they presented a remarkable contrast in appearance with the adjoining uncovered plants, which were densely covered with capsules. In 1863 a protected long-styled plant produced only five poor capsules; two mid-styled plants produced the same number; and two short-styled plants between them produced only one: these capsules contained very few seed; yet the plants were fully productive when artificially fertilized under the net. In a state of nature the flowers are incessantly visited for their nectar by hive- and humble-bees and various Diptera. The nectar is secreted all round the base of the ovarium; but a passage is formed along the upper and inner side of the calyx by the lateral deflection (not represented in the diagram) of the basal portions of the filaments; so that insects invariably alight on the upper side of the flowers, on the projecting stamens and pistil, and insert their probosces along the upper inner margin of the calyx. We can now see why the ends of the stamens with their anthers, and the ends of the pistils with their stigma, are a little upturned, in order that they may brush against the lower hairy surfaces of the insects' bodies. The short stamens which lie enclosed within the calyx of the long- and mid-styled forms can be touched only by the proboscis and the narrow chin of the sucking bee; hence they have their ends more upturned, and they are graduated in length, so as to fall into a narrow file, three deep, sure to be raked by the thin intruding proboscis. The anthers of the longer stamens stand laterally further apart and are more nearly of the same length, for they have to brush against the whole breadth of the insect's body. I may here incidentally remark, that in very many flowers the pistil, or the stamens, or both, are rectangularly bent to one side of the flower: this bending may be permanent, as with Lythrum and many others, or may be effected (as in Dictamnus fraxinella and many others) by a temporary movement which occurs in the stamens when the anthers dehisce, and in the pistil when the stigma is mature; but these two movements are by no means always contemporaneous in the same flower. Now I have found no exception to the rule, that when the stamens and pistil are bent, the bending is exactly to that side of the flower which secretes nectar (even though there be a rudimentary nectary of large size on the opposite side, as in some species of Corydalis); or, when nectar is secreted on all
sides, to that side where the structure of the flowers allows the easiest access to it, as in Lythrum, Papilionaceous flowers, and many others. The rule consequently is that when the pistil and stamens are bent, the stigma and anthers are brought into the pathway towards the nectary. There are a few cases which seem to be exceptions, but they are not so in truth: for instance, in the Gloriosa lily, the stigma of the grotesque and rectangularly bent pistil is brought, not into the pathway from the open air towards the nectar-secreting recesses of the flower, but into the circular route from one nectary to the other; in Scrophularia aquatica the pistil is bent downwards from the mouth of the flower, but it thus strikes the pollen-dusted breasts of the wasps which habitually visit these ill-scented blooms. In the above rule we see one more instance of the supreme dominating power of insects over all the minor structural details of flowers, especially of those which have irregular corollas. Flowers which are fertilized by the wind must of course be excepted, but I do not know of a single instance of an irregular flower which is fertilized or crossed by this means.
I have delayed too long on these points, but I must allude to one other. We have seen that the three pistils of different lengths have each two half-dozen sets of stamens of corresponding length. When bees suck the flowers, the longest stamens, bearing the green pollen, rub against the abdomen and the interior sides of the posterior legs, as does likewise the stigma of the long-styled form. The stamens of middle length and the stigma of the mid-styled form rub against the under side of the thorax and between the front pair of legs. The shortest stamens and the stigma of the short-styled form must rub against the proboscis and chin; for the bees in sucking insert only the front of their heads into the calyx. On catching bees, I observed much green pollen on the inner sides of the hind legs and on the abdomen, and much yellow pollen on the under side of the thorax. There was also pollen on the chin, and, it may be presumed, on the proboscis, but this was difficult to observe. I had, however, independent proof that pollen is carried on the proboscis; for in a protected short-styled plant (which produced only two capsules) one small branch was accidentally left during many days pressing against the fine net, and bees were seen inserting their probosces through the meshes, and in consequence numerous capsules were formed on this one small branch. From these several facts it follows that insects would chiefly carry to the
stigma of each form pollen from the stamens of corresponding length; and we shall presently see the importance of this adaptation. It must not, however, be supposed that the bees do not get more or less dusted all over with the several kinds of pollen; they certainly do, as could be seen with the green pollen from the longest stamens. Moreover, a case will presently be given of a long-styled plant which grew absolutely by itself, and produced an abundance of capsules, which must have been fertilized by its own two kinds of pollen; but these capsules contained a very poor average of seed. Hence insects, and chiefly bees, act both as general carriers of pollen, and as special carriers of the right kind*.
Variability.—Before passing on to more important topics, I must say a few words on this head. Wirtgen remarks† on the variability in the branching of the stem, in the length of the bracteæ, size of the petals, and in several other respects. The plants now growing in my garden have their leaves arranged oppositely, alternately, and in whorls of three, and differ greatly in shape. The stems of the plants bearing leaves in whorls are hexagonal; those of the other plants are quadrangular. But we are concerned only with the reproductive organs: the upward bending of the pistil is variable, and in a remarkable degree in the short-styled form, in which it is sometimes straight, sometimes slightly curved, but generally upturned at right angles. The stigma of the long-styled pistil frequently has longer papillæ or is rougher than that of the mid-styled, and this than that of the short-styled form; but this character, though fixed and uniform with the two forms of Primula, is here variable, and I have
* In my paper on the two forms of Primula (Journal Proc. Linn. Soc. 1862, p. 85) [Darwin 1862] I stated that I had only occasionally seen humble-bes [sic] sucking the flowers of the Cowslip (P. veris). Since then I have had some beds in my garden containing nearly 700 plants, and these were incessantly visited by Bombus hortorum and B. muscorum. I caught some of these bees, and I found (as I had anticipated in my paper, p. 86) that a vast majority of the pollen-grains which adhered to the base of the proboscis were large-sized and had come from the long stamens of the short-styled form, and were thus placed ready to fertilize the stigma of the long-styled form. On the other hand, on the middle, and near the tip of the proboscis, a very large proportion of the pollen-grains were of the small size, and had come from the short stamens of the long-styled form. My son caught, also, a moth (Cucullia verbasci) hovering over the bed, and I found on its proboscis a similar distribution of the two kinds of pollen-grains. I give these facts as a further illustration of the importance of the relative lengths of the stamens and pistil.
† Verhand. des naturhist. Vereins, 5. Jahrgang, 1848, S. 11, 13.
seen mid-styled stigmas rougher than those of the long-styled. The degree to which the longer and middle stamens are graduated in length and are upturned at their ends is variable; sometimes all are equal. The colour of the green pollen in the long stamens is variable*, and is sometimes pale greenish yellow; in one short-styled plant it was almost white. The grains vary a little in size: I examined one short-styled plant with the grains above the average size; and I have seen a long-styled plant with undistinguishable grains from the longer and shorter anthers. We have here considerable fluctuations of character; and if any of these slight structural differences were of direct service to the plant, or were correlated with useful functional differences, we can perceive that the species is just in that state in which natural selection might readily do much for its modification.
To return to our proper subject—we see that there are three kinds of females and three kinds of males, each kind of the latter being borne by half-dozens on two of the three forms. It remains to discover whether these several sexes or sexual organs differ from each other in function. Nothing brings more prominently forward the complexity of the reproductive system of this extraordinary plant, than the necessity, in order to ascertain the above fact, of artificially making eighteen distinct unions. Thus the long-styled form had to be fertilized with pollen from its own two distinct kinds of anthers, from the two in the mid-styled, and from the two in the short-styled form. The same process had to be repeated with both the mid- and short-styled forms. It might have been thought sufficient to have tried on each stigma the green pollen, for instance, from either the mid- or short-styled longer stamens, and not from both; but the result proves that this would have been insufficient, and that it was necessary to try all six kinds of pollen on each stigma. As in artificial fertilizations there will always be some failures, it would have been advisable to have
* Lagerstrœmia Indica, one of the Lythraceæ, is strangely variable in its stamens—I presume in part due to its growth in a hothouse. The most perfect flowers produced with me five very long stamens with thick flesh-coloured filaments and green pollen, and from nineteen to twenty-nine short stamens with yellow pollen; but many flowers produced only one, two, three, or four long stamens with green pollen, which in some of the anthers was wholly replaced by yellow pollen; one anther offered the singular case of half, or one cell being filled with bright green, and the other cell with bright yellow pollen. One petal had a furrow near its base, which contained pollen. According to analogy with Lythrum, this species would produce three forms; if so, the above plant was a mid-styled form: it was quite sterile with its own two kinds of pollen.
repeated each of the eighteen unions a score of times; but the labour would have been too great; as it was, I made 223 artificial unions; i. e., I fertilized, on an average, above a dozen flowers in the eighteen different methods. Each flower was castrated; the adjoining buds had to be removed, that the marking-thread, wool, &c. might be safely secured; and after each fertilization the stigma had to be examined with a lens to see that there was sufficient pollen. Plants of all three forms were protected during two years by large nets on a framework; two plants were used during one or both years, in order to avoid any individual peculiarity in any one plant. As soon as the flowers withered, the nets were removed; and in the autumn the capsules were daily inspected; when the seeds were ripe they were counted under the microscope. I have given these details that confidence may be placed in the following Tables, and as some excuse for two blunders which, I believe, I made. These blunders are referred to, with their probable causes, in two notes to the Tables; the erroneous numbers, however, are entered in the Tables, that it may not be supposed that I have in any one instance tampered with the results.
A few words explanatory of the three Tables must be given. Each is devoted to one form, and is divided into six compartments. The two upper ones in each table give the product of good seed from the application of pollen from the two sets of stamens which correspond in length with the pistil of that form. The two next lower compartments show the result of pollen from the other two sets of stamens, which do not correspond in length with the pistil, and which are borne by the same two forms. The two lowest compartments show the result of the application of each form's own two kinds of pollen. The term "own pollen," used here and in the Tables, does not mean pollen from the flower to be fertilized—for this was never used—but from another flower on the same plant, or more commonly from a distinct plant of the same form. In the result given, "0" generally means that no capsule was produced, or that the capsule contained no good seed. In some part of each row of figures in each compartment, a short horizontal line may be seen; the unions above this line were made in 1862, and below it in 1863. It is of importance to observe this, as it shows that the same general result ensued in two successive years; but more especially because 1863 was a very hot and dry season, and the plants had occasionally to be watered. This did not prevent the full complement of seed being produced from
the more fertile unions; but it rendered the less fertile unions even more sterile than they otherwise would have been. I have seen striking instances of this same fact in making homomorphic and heteromorphic unions in Primula*; and it is well known that the conditions of life must be highly favourable to give any chance of producing hybrids from species which cross with difficulty.
Table I.—LONG-STYLED FORM.
|13 flowers fertilized by the longer stamens of the mid-styled. These stamens equal in length the pistil of the long-styled.
|13 flowers fertilized by the longer stamens of the short-styled. These stamens equal in length the pistil of the long-styled.
|Product of good seed in each capsule.
|Product of good seed in each capsule.
|96 poor seed.
|38 per cent. of these flowers yielded capsules. Each capsule contained, on an average, 51.2 seed.
|84 per cent. of these flowers yielded capsules. Each capsule contained, on an average, 107.3 seed.
* In the spring of 1862 I crossed forty Cowslip flowers (P. veris) heteromorphically and homomorphically. The plants were accidentally exposed in the greenhouse to too hot a sun, and a number of umbels perished. Some, however, remained in moderately good health, and on these there were twelve flowers which had been fertilized heteromorphically and eleven which had been fertilized homomorphically. The twelve heteromorphic unions yielded seven fine capsules, containing on an average 57.3 good seed. Now mark the difference: the eleven homomorphic unions yielded only two capsules, of which one contained 39 seeds, but so poor, that I do not suppose one would have germinated, and the other only 17 fairly good seed. It would be superfluous to give any more details on this experiment, or on some which I made at the same time on P. Sinensis, after the appearance of Mr. John Scott's admirable paper1 on the various dimorphic species of Primula, in which he confirms my former results, and adds many original and valuable observations. Dr. Hildebrand2 has also (Botanische Zeitung, 1864, Jan. 1, S. 3) confirmed my general results with respect to P. Sinensis, and has corrected an error into which in some unaccountable manner I fell, namely, that the pollen-grains from the long- and short-styled forms were of the same size. Dr. Hildebrand has added a series of new and important experiments, for he fertilized homomorphically a number of flowers with pollen from the same form, and likewise from the same individual flower. These latter he found were thus rendered rather more sterile. This experiment, I believe, has never been systematically tried before.
1 Scott 1864.
2 Friedrich Hermann Gustav Hildebrand (1835-1915), German professor of botany, University of Freiburg, 1868-1907. Hildebrand 1864.
Table I.—LONG-STYLED FORM (continued).
|14 flowers fertilized by the short stamens of the mid-styled.
|12 flowers fertilized by the shorter stamens of the short-styled.
Too sterile for any average.
Too sterile for any average.
|15 flowers fertilized by own longer stamens.
|15 flowers fertilized by own shorter stamens.
Too sterile for any average.
Too sterile for any average.
I fertilized a considerable number of flowers with pollen, taken by a camel's-hair brush, from both the long and short stamens of their own (long-styled) form; but I did not examine with a lens (as I did in the cases in the Tables) whether sufficient pollen had been placed on the stigma: only 5 capsules were produced, and these yielded on an average 14.5 seed. In 1863 I tried a much better experiment: a long-styled plant was grown by itself, miles away from any other plant, so that its stigmas could have received only the two kinds of pollen proper to this form. The flowers were incessantly visited by bees, so that the stigmas must have received on the most favourable days, and at the most favourable hours, successive applications of pollen: all who have crossed plants know that this highly favours fertilization. This plant produced an abundant crop of capsules; I took by chance 20, and these (excluding one poor one) contained seed as below:—
This gives an average of 21.5 seed per capsule; and as we know that this form, when standing near plants of the other two forms and fertilized by insects, produces an average of 93.1 seed per capsule, we see that the long-styled form fertilized by its own two pollens yields only between one-fourth and one-fifth of the full number of seed. I have spoken as if this plant had received both its own kinds of pollen, and this is, of course, possible; but, from the enclosed position of the shorter stamens, it is much more probable that the stigma received almost exclusively the pollen from its own longer stamens.
Table II.—MID-STYLED FORM.
|12 flowers fertilized by the longer stamens of the long-styled. These stamens equal in length the pistil of the mid-styled.
|12 flowers fertilized by the shorter stamens of the short-styled. These stamens equal in length the pistil of the mid-styled.
|Product of good seed in each capsule.
|Product of good seed in each capsule.
|92 per cent. of the flowers (probably 100 per cent.) yielded capsules. Each capsule contained, on an average, 127.3 seed.
|100 per cent. of the flowers yielded capsules. Each capsule contained, on an average, 108.0 seed; or, excluding capsules with less than 20 seed, the average is 116.7 seed.
|13 flowers fertilized by the short stamens of the long-styled.
|15 flowers fertilized by the longer stamens of the short-styled.
|85 {seed small and poor.
| 6
|54 per cent. of the flowers yielded capsules. Each capsule contained, on an average, 47.4 seed; or, excluding capsules with less than 20 seed, the average is 60.2 seed.
|93 per cent. of the flowers yielded capsules. Each capsule contained, on an average, 69.5 seed; or, excluding capsules with less than 20 seed, the average is 102.8.
|12 flowers fertilized by own longer stamens.
|12 flowers fertilized by own shorter stamens.
|Excluding the capsule with 136 seed, 25 per cent. of the flowers yielded capsules, and each capsule contained, on an average, 54.6 seed; or, excluding capsules with less than 20 seed, the average is 77.5.
|0 per cent. of the flowers yielded capsules.
* I have hardly any doubt that this result of 136 seed is due to a gross error. The flowers to be fertilized by their own longer stamens were first marked by
I fertilized a considerable number of flowers with pollen, taken by a camel's-hair brush, from both the long and short stamens of their own (short-styled) form; but I did not examine with a lens (as I did in the cases in the Tables) whether sufficient pollen had been placed on the stigma: only 5 capsules were produced, and these yielded, on an average, 11.0 seed.
Table III.—SHORT-STYLED FORM.
|12 flowers fertilized by the shorter stamens of the long-styled. These stamens equal in length the pistil of the short-styled.
|13 flowers fertilized by the shorter stamens of the mid-styled. These stamens equal in length the pistil of the short-styled.
|83 per cent. of the flowers yielded capsules. Each capsule contained, on an average, 81.3 seed.
|61 per cent. of the flowers yielded capsules. Each capsule contained, on an average, 64.6 seed.
|10 flowers fertilized by the longer stamens of the long-styled.
|10 flowers fertilized by the longer stamens of the mid-styled.
|Too sterile for any average.
|Too sterile for any average.
"white thread," and those by the longer stamens of the long-styled form by "white silk;" a flower fertilized in the latter manner would have yielded about 136 seed, and it may be observed that one such pod is missing, viz., at the bottom of compartment 1. Therefore I have hardly any doubt that I fertilized a flower marked with "white thread," as if it had been marked with "white silk." With respect to the capsule which yielded 92 seed, in the same column with that which yielded 136, I do not know what to think. I endeavoured to prevent pollen dropping from an upper to any lower flower, and I tried to remember to wipe the pincers carefully after each fertilization; but in making eighteen different crosses, sometimes on windy days, and pestered by bees and flies buzzing about, some few errors could hardly be avoided. One day I had to keep a third man by me all the time to prevent the bees visiting the uncovered plants, for in a few seconds' time they might have done irreparable mischief. It was also extremely difficult to exclude minute Diptera from the net. In 1862 I made the great mistake of placing a mid-styled and long-styled under the same huge net: in 1863 I avoided this error.
Table III.—SHORT-STYLED FORM (continued).
|10 flowers fertilized by own longer stamens.
|10 flowers fertilized by own shorter stamens.
|Too sterile for any average.
|Too sterile for any average.
I fertilized a number of flowers without particular care with their own two pollens, but they did not produce a single capsule; the position of the stigma within the calyx renders the fertilization without some care difficult.
Summary of the three preceding Tables.
Long-styled form.—Twenty-six flowers fertilized by the stamens of corresponding length, borne by the mid- and short-styled forms, yielded 61.5 per cent. of capsules, which contained, on an average, 89.7 seed.
Twenty-six flowers fertilized by the other and shorter stamens of the mid- and short-styled forms yielded only two very poor capsules.
Thirty flowers fertilized by this form's own two sets of stamens yielded only eight very poor capsules; but flowers well fertilized by bees by one or both of their own kinds of pollen produced numerous capsules containing, on an average, 21.5 seed.
Short-styled form.—Twenty-five flowers fertilized by the stamens of corresponding length, borne by the long- and mid-styled forms, yielded 72 per cent. of capsules, which (excluding one capsule with only nine seeds) contained, on an average, 70.8 seed.
Twenty flowers fertilized by the longer stamens of the long- and mid-styled forms yielded only two very poor capsules.
Twenty flowers fertilized by both their own two sets of stamens yielded only two poor (or perhaps three) capsules.
Mid-styled form.—Twenty-four flowers fertilized by the stamens of corresponding length, borne by the long- and short-styled forms, yielded 96 (probably 100) per cent. of capsules, which con-
* I suspect that, by mistake, I fertilized this flower with the pollen of the shorter stamens of the long-styled form, and it would then have yielded about 64 seed. Flowers to be thus fertilized were marked with black silk; those with the pollen of the shorter stamens of the short-styled with black thread; and thus, I suspect, the mistake arose.
tained (excluding one capsule with 12 seed), on an average, 117.2 seed.
Fifteen flowers fertilized by the longer stamens of the short-styled form yielded 93 per cent. of capsules, which (excluding four capsules with less than 20 seed) contained, on an average, 102.8 seed.
Thirteen flowers fertilized by the shorter stamens of the long-styled form yielded 54 per cent. of capsules, which capsules (excluding one with 19 seed) contained, on an average, 60.2 seed.
Twelve flowers fertilized by own longer stamens yielded 25 per cent. of capsules, which (excluding one with 9 seed) contained, on an average, 77.5 seed.
Twelve flowers fertilized by own shorter stamens yielded not a single capsule.
Considering the three Tables and this summary, we may safely draw the following conclusions. First, that, as in structure so in function, there are three females or female organs: this is manifest; for when all three receive the very same pollen, they are acted on most differently. So conversely with the thirty-six stamens, we know that they consist of three separate sets of a dozen each, differing in various respects; and in function the pollen of these three sets when applied to one and the same stigma acts most differently, as a glance at the Tables proves. But we shall presently see that the action of the pollen of the whole dozen longest and of the whole dozen shortest stamens is not identical.
Secondly, we see that only the longest stamens fully fertilize the longest pistil, the middle stamens the middle pistil, and the shortest stamens the shortest pistil. And now we can comprehend the meaning of the almost exact correspondence in length between the pistil of each form and the two half-dozen sets of stamens borne by the two other forms; for the stigma of each form is thus rubbed against the same spot of the insect's body, which becomes most charged with the proper pollen. In all three forms, the female organ is but feebly, or not at all, acted on by its own two kinds of pollen. In my papers on the dimorphism of Primula and Linum, I used the terms "heteromorphic" for the fully fertile unions between the female element of the one form and the male element of the other, and "homomorphic" for the less fertile or quite sterile unions between the female and male elements of the same form. The principle involved in these terms holds good with Lythrum, but is insufficient; for though in each of the three forms the fertile unions are all heteromorphic, the ap-
propriate pollen coming from the stamens of corresponding length borne by the other two forms, and though the homomorphic unions of the females with their own two sets of males are always more or less sterile, there remain in each case two other sterile unions, not included in these two terms. Hence it will be found convenient to designate the two unions of each female with the two sets of stamens of corresponding length, which are fully fertile, as legitimate unions, and the four other, more or less sterile, unions of each female with the four other sets of stamens as illegitimate unions. Consequently, of the eighteen possible unions between the three forms, six are legitimate and twelve are illegitimate.
Another and curious conclusion cannot be considered as proved, but is rendered highly probable, by the Tables. The unions of the pistils and stamens of equal length are alone fully fertile. Now with the several illegitimate unions it will be found that the greater the inequality in length between the pistil and stamens, the greater the sterility of the result. There is no exception to this rule. Thus, with the long-styled form, its own shorter stamens are far less equal in length to the pistil than its own longer stamens; and the capsules fertilized by the pollen of the shorter stamens yielded fewer seeds: the same comparative result follows from the use of the pollen of the shorter stamens of the mid-styled form, which are much shorter than the shorter stamens of the short-styled (see diagram), and therefore less equal in length to the long-styled pistil. We shall see exactly the same result if we look to the four illegitimate unions under the mid- and short-styled forms. Certainly the difference in sterility in these several cases is very slight, but the sterility always increases with the increasing inequality of length between the pistil and the stamens which are used. Therefore I believe in the above rule; but a vast number of artificial unions would be requisite to prove it. If the rule be true, we must look at it as an incidental and useless result of the gradational changes through which this species has passed in arriving at its present condition. On the other hand, the correspondence in length between the pistil of each form and those stamens which alone give full fertility is clearly of service to the species, and is probably the result of direct adaptation.
Some of the illegitimate unions yielded, as may be seen in the Tables, during neither year a single seed; but, judging from the case of the long-styled plant, it is probable, if such unions could be effected repeatedly under the most favourable conditions, some few seeds would be produced. Anyhow, I can state that in all
the eighteen possible unions the pollen-tubes penetrated, after eighteen hours, the stigma. I have reason to believe that the offspring from the illegitimate unions present some singular characteristics; but until my observations on this head are repeated, I must be silent. At first I thought that perhaps two kinds of pollen placed together on the same stigma would give more fertility than any one kind; but we have seen that this is not the case with each form's own two kinds of pollen; nor is it probable in any case, as I occasionally got, by the use of single kinds of pollen, fully as many seed as I have seen in a capsule naturally fertilized. Moreover the proper pollen from a single anther is more than sufficient to fully fertilize each stigma; hence, in this as in so many other cases, at least twelve times as much of each kind of pollen is produced as is necessary to ensure full fertilization. From the dusted condition of the whole body of those bees which I caught on these flowers, it is probable that some pollen of all kinds is deposited on each stigma; but there can hardly be a doubt that the pollen of the stamens of corresponding length will be prepotent and will wholly obliterate any effect from the other kinds of pollen, even if previously deposited on the stigma. I infer this partly from the fact ascertained by Gärtner that each species' own pollen is so prepotent over that of any other species, that if put on the stigma many hours subsequently, it will entirely obliterate the action of the foreign pollen. But I draw the above inference especially from the following experiment: I fertilized homomorphically or illegitimately some long-styled Cowslip flowers (Primula veris) with their own pollen, and exactly twenty-four hours subsequently I fertilized these same stigmas heteromorphically or legitimately with pollen from a short-styled dark-red Polyanthus. I must premise that I have raised many seedlings from crossed Cowslips and Polyanthus, and know their peculiar appearance; and I further know, by the test of the fertility of the mongrels inter se, and with both parent forms, that the Polyanthus is a variety of the Cowslip, and not of the Primrose (P. vulgaris) as some authors have supposed. Now from the long-styled Cowslip twice fertilized in the manner explained, I raised twenty-nine seedlings, and every one of them had flowers coloured more or less red; so that the heteromorphic Polyanthus-pollen wholly obliterated the influence of the homomorphic pure Cowslip-pollen, which had been placed on the stigmas twenty-four hours previously, and not a single pure Cowslip was produced.
The last conclusion which may be deduced from the Tables, even from a glance at them, is that the mid-styled form differs from both the others in its much higher capacity for fertilization. Not only did the twenty-four flowers fertilized by the stamens of corresponding lengths, all, or all but one, yield capsules rich in seed; but of the other four illegitimate unions, that by the longer stamens of the short-styled form was highly fertile, though less than in the two legitimate unions, and that by the short stamens of the long-styled form was fertile to a considerable degree; the two unions with this form's own pollen were sterile, but in different degrees. So that the mid-styled form, when fertilized by the six kinds of pollen, evinces five different grades of fertility. By comparing compartments 3 and 6 in Table II. we learn a remarkable fact, namely, that though the pollen from the short stamens of the long-styled and from this form's own (mid-styled) short stamens, used in these two unions, is identical in all respects, yet that its action is widely different; in the one case above half the fertilized flowers yielded capsules containing a fair number of seed; in the other case not one single capsule was produced. So, again, the green, large-grained pollen from the long stamens of the short-styled and from this form's own (mid-styled) long stamens is identical in all respects, but its action, as may be seen in compartments 4 and 5, is widely different. In both these cases the difference in action is so plain that it cannot be mistaken, but it can be corroborated. If we look to Table III., to the legitimate action of the short stamens of the long- and mid-styled forms on the pistil of the short-styled form, we again see a similar but slighter difference, the pollen of the short stamens of the mid-styled form yielding a smaller average of seed during the two years of 1862 and 1863 than that from the short stamens of the long-styled form. Again, if we look to Table I., to the legitimate action of the green pollen of the two sets of long stamens, we shall find exactly the same result, viz. that the pollen of the long stamens of the mid-styled form yielded during both years fewer seeds than that from the long stamens from the short-styled form. Hence it is certain that the two kinds of pollen produced by the mid-styled form are less potent than the similar pollens produced by the corresponding stamens of the two other forms.
When we see that the capsules of the mid-styled form yield a considerably larger average number of seed than those of the other two forms,—when we see how surely the flowers are fertilized in the legitimate unions, and how much more productive
the illegitimate unions are than those of the other two forms, we are led to consider the mid-styled form as eminently feminine in its nature. And although it is impossible to consider as rudimentary or aborted the two perfectly developed sets of stamens of the mid-styled form which produce an abundance of perfectly well-developed pollen, yet we can hardly avoid connecting, as balanced, the higher efficiency of the female organ with the lesser potency of the two mid-styled pollens.
Finally, it is proved by the Tables that Lythrum salicaria habitually produces or consists of three females different in structure and widely different in function; that it produces or consists of three sets of males widely different in structure and function; and that two of the three sets of males are subdivided into subgroups of half a dozen each, differing in a marked manner in potency, so that regularly five kinds of pollen are elaborated by this one species of Lythrum.
Lythrum Græfferi.—I must now say a few words about some of the other species of the genus. I have examined numerous dried flowers of L. Græfferi, each from a separate plant, kindly sent me from Kew. This species, like L. salicaria, is trimorphic, and the three forms apparently occur in about equal numbers. In the long-styled form the pistil projects about one-third of the length of the calyx beyond its mouth, and is therefore shorter than in L. salicaria; the globose and hirsute stigma is larger than that of the other two forms; the longer stamens, which are graduated in length, have their anthers standing just above and just beneath the mouth of the calyx; the half-dozen shorter stamens rise rather above the middle of the calyx. In the mid-styled form the stigma projects just above the mouth of the calyx, and stands almost on a level with the longer stamens of the previous form; its own longer stamens project well above the mouth of the calyx and stand a little above the level of the stigma of the long-styled form; the shorter stamens correspond in all respects with the shorter ones in the previous form. In the short-styled form the stigma of the pistil is nearly on a level with the anthers of the shorter stamens in the two preceding forms; and the longer stamens correspond with the longer stamens of the mid-styled form, and the shorter stamens with the longer stamens of the long-styled form. In short, there is a close general correspondence in structure between this species and L. salicaria, but with some differences in the proportional lengths of the parts. Nevertheless the fact of each of the three pistils having two sets
of stamens, borne by the two other forms, of corresponding lengths, comes out conspicuously. In the mid-styled form the distended pollen-grains from the longer stamens had nearly double the diameter of those from the shorter stamens; so that there is a greater difference in this respect than in L. salicaria. In the long-styled form, also, the difference in diameter between the pollen-grains of the longer and shorter stamens was plainer than in L. salicaria. These comparisons, however, must be received with caution, as they were made on specimens long kept in a dried condition.
Lythrum thymifolia.—This form, according to Vaucher*, is dimorphic like Primula, and therefore presents only two forms. I received two dried flowers from Kew, which presented two forms: in the one form the stigma projected far beyond the calyx, in the other it was included within the calyx; in this latter form the style was only one-fourth of the length of the style of the other form. There are only six stamens; these are somewhat graduated in length, and in the short-styled form the anthers stand a little above the stigma, but yet the stamens by no means equal in length the pistil of the long-styled form; in the long-styled form the stamens are rather shorter than in the other form. These six stamens alternate with the petals, and correspond homologically with the longer stamens of L. salicaria and L. Græfferi. As there are only six stamens, it is scarcely possible that this species can be trimorphic.
Lythrum hyssopifolia.—This species is said by Vaucher, but I believe erroneously, to be dimorphic. I have examined dried flowers from twenty-two separate plants from various localities, kindly sent to me by Mr. Hewett C. Watson,1 Prof. Babington, and others. These were all essentially alike. Hence the species cannot be dimorphic. The pistil varies somewhat in length, but when unusually long the stamens are likewise generally long; in the bud the stamens are short: perhaps these circumstances deceived Vaucher. There are from six to nine stamens, graduated in length; the stamens which are variable in being present or absent correspond with the six shorter stamens of L. salicaria and with the six which are absent in L. thymifolia. The stigma is included within the calyx, and stands in the midst of the anthers, and would generally be fertilized by them; but as the stigma and anthers are upturned, and as, according to Vaucher, there is a passage left in the upper side of the flower to the nectary, there can hardly be a doubt that
* Hist. Phys. des Plantes d'Europe, tom. ii. (1841) pp. 369, 371.
1 Hewett Cottrell Watson (1804-1881), botanist and phrenologist who specialized in the distribution of British plants.
the flowers are visited by insects, which would occasionally bring pollen from other flowers of the same or of any adjoining plant, as surely as occurs with the short-styled L. salicaria, of which the pistil and corresponding stamens closely resemble those of L. hyssopifolia. According to Vaucher and Lecoq*, this species, which is an annual, generally grows almost solitarily, whereas the three preceding species are social; and this alone would almost have convinced me that L. hyssopifolia cannot be dimorphic, as such plants cannot habitually live by themselves any better than one sex of a diœcious species.
Nesæa verticillata.—I raised a number of plants from seed sent me by Professor Asa Gray, and they presented three forms. These differed from each other in the proportional lengths of their organs of fructification and in all respects in very nearly the same way as the three forms of Lythrum Græfferi. The green pollen-grains from the longest stamens, measured along their greater axis and not distended with water, were 13/7000 of an inch in length; those from the stamens of middle length 9-10/7000, and those from the shortest stamens 8-9/7000 of an inch.
We have seen that the genus Lythrum affords trimorphic, dimorphic, and monomorphic species.
The inquiry naturally arises, why do these species differ so remarkably in their sexual relations? of what service can reciprocal dimorphism or trimorphism be to certain species, whilst other species of the same genus present, like the great majority of plants, only one form? I have elsewhere given too briefly† the
* Géograph. Bot. de l'Europe, tom. vi. (1857) p. 157.
† 'Origin of Species,' 3rd edit., p. 101. Hugo von Mohl1 has recently (Bot. Zeitung, 1863, S. 309, 321), in a most interesting paper, advanced the case of the minute, imperfectly developed, closed and self-fertile flowers borne by Viola, Oxalis, Impatiens, Campanula, &c., as an argument against my doctrine that no species is self-fertilized for perpetuity. I may state that in the spring of 1862 I examined some of these flowers, and saw, though less thoroughly, all that H. von Mohl has so well described. I can add only one remark, which I believe is correct, that in V. canina there is an open channel for the pollen-tubes from the extremity of the stigma to the ovarium; for I gently pressed a minute bubble of air repeatedly backwards and forwards from end to end. Though the imperfectly developed and the perfect flowers are so different in structure, it is a rather curious case of correlation, that in the double purple Violet (V. odorata) the minute imperfect flowers are double to the very core, so that a section appears like the head of a cabbage when cut through. There can be, as von Mohl asserts, no doubt that these flowers are always self-fertilized; they are moreover specially adapted for this end, as may be seen in the remarkable difference in the shape of the pistil in V. canina (and in a less degree in V. hirta and the single V. odorata) as compared with that of the perfect flower;
1 Hugo von Mohl (1805-1872), German botanist and physiologist who co-founded the Botanische Zeitung, 1843. Mohl 1863.
general grounds of my belief that with all organic beings distinct individuals at least occasionally cross together, and reciprocal dimorphism is plainly one most efficient means for ensuring this result.
and in the pollen-tubes which proceed from the grains within the anthers in V. canina, and from within the lower anthers of Oxalis acetosella, having the wonderful power of directing their course to the stigma. If these plants had produced the minute closed flowers alone, the proof would have been perfect that they could never have crossed with other individuals. I am aware that in some of these cases it has been stated that the perfect flowers never produce any seed; as far as Amphicarpæa is concerned, I hear from Professor Asa Gray that the Petaliferous flowers certainly sometimes yield seed. The completely enclosed flowers of that curious grass, the Leersia oryzoides, as described by M. Duval-Jouve1 (Bull. Soc. Bot. de France, tom. x. 1863, p. 194), apparently offer the best case of perpetual self-fertilization; for when perfect flowers are protruded from the culms, they are, as far as is yet known, always sterile. In a number of plants kept by me in pots in water, not one single perfect flower has protruded, but the enclosed flowers produced plenty of seed. Without wishing to throw any doubt on M. Duval-Jouve's excellent observations, I may add that with the enclosed flowers borne by my plants, the act of fertilization, that is, the penetration of the stigma by the pollen-tubes, took place in the air and not in fluid within the glumes. With the exception of the Leersia, as the case now stands, I cannot see how the production of the small, imperfect flowers invalidates my doctrine that no species is perpetually self-fertilized, more than the multiplication of many plants by bulbs, stolons, &c. As I observe that the production of seed by the perfect flowers of Viola is spoken of as something capricious and accidental, I may state that, although it varies much in different years, it depends exclusively on the visits of bees; I ascertained this by marking many flowers thus visited, and finding that they produced capsules, and by covering up many flowers which (excepting a few that I artificially fertilized) did not, when thus protected, produce a single capsule. After bees have visited these flowers, the pollen may be seen scattered on the papillæ and on the stigma itself, and they can hardly fail thus to cross distinct individuals. These remarks apply to V. canina, hirta, and odorata; with V. tricolor the case is somewhat different; but I must not enlarge any more on this subject. The production by so many plants of perfect and expanded, as well as of imperfect and closed flowers, seems to me to throw much light on many points; it shows how extraordinarily little pollen is necessary for full fertilization, for I ascertained with V. canina that the perfect and imperfect flowers (the latter producing so few pollen-grains) yielded the same average number of seeds; it shows us that fertilization can be perfected in closed flowers; it shows us that large, highly coloured petals, perfume, and the secretion of nectar are by no means indispensable for this act, even in those species which properly possess these characters. It seems to me that the necessity of an occasional cross with a distinct individual of the same species explains the universal presence of at least some expanded flowers, at the expense of injury from rain and the loss of much pollen by innumerable pollen-robbing insects; it explains the enormous superfluity of pollen from its liability to loss from these causes and during conveyance from flower to flower; it explains the use of a gaily coloured corolla, perfume, and nectar, namely, to attract insects, except in those comparatively few cases in which wind is the agent, and in these the last-named attributes are deficient.
1 Joseph Duval-Jouve (1810-1883), French botanist.
This result would appear to be one of high importance, for with dimorphic plants it is ensured at the risk of occasional sterility; not only is the pollen of each plant useless or nearly useless to that individual, but so is the pollen of all the plants of the same form, that is, of half the total number of individual plants. In that extensive class of plants called by C. K. Sprengel dichogams, in which the pollen of each flower is shed before its own stigma is ready, or in which the stigma (though this case occurs more rarely) is mature before the flower's own pollen is ready sterility can hardly fail to be the occasional result; and it would be the inevitable result with both dichogamous and reciprocally dimorphic flowers unless pollen were carried by insects (and in some few species by the wind) from one flower or plant to the other. As with reciprocal dimorphism so with dichogamy, within the same genus some of the species are and some are not thus characterized. Again, in the same genus, as in that of Trifolium, some species absolutely require insect-aid to produce seed, others are fertile without any such aid; now when insects are requisite for fertilization, pollen will generally be carried from one flower to the other. We thus see, by means of reciprocal dimorphism, of dichogamy, and of insect-aid, that some species require, or at least receive, incessant crosses with other individuals of the same species; whereas other species of the same genera can be, and probably are often fertilized during long periods by the pollen of their own flowers. Why this wide difference in the frequency of crosses should occur we are profoundly ignorant. I will only further remark on this head, that it would be a great mistake to suppose that many flowers, which are neither reciprocally dimorphic nor dichogamous, nor require insect-aid for their fertilization, nor show any particular adaptation in their structure for the visits of insects, are not habitually crossed with the pollen of other individuals; this occurs, for instance, habitually with cabbages, radishes, and onions, which nevertheless are perfectly fertile (as I know by trial) with their own pollen without aid of any kind.
But it may be further asked, granting that reciprocal dimorphism is of service by ensuring at each generation a cross (but I am far from pretending that it may not have some additional unknown signification), why did not dimorphism suffice for L. salicaria and Græfferi? why were they rendered reciprocally trimorphic, entailing such complicated sexual relations? We cannot answer, except perhaps so far:—if we suppose two plants of
the L. salicaria to grow by themselves, then if the species were dimorphic it would only be an equal chance in favour of the two turning out different forms and consequently both being fertile; but as the species is trimorphic and each form can fertilize the two other forms, it is two to one in favour of the two turning out different forms and being consequently both fertile. We thus see how reciprocal trimorphism must be an advantage; and probably it would be more advantageous to this Lythrum, which commonly grows in almost a single row along the banks of streams, than it would be to Primroses or Cowslips which have neighbours on all sides. But even if trimorphism effected no good beyond that gained by dimorphism, we ought not to feel much surprised at its occurrence, for we continually see throughout nature the same end gained by the most complicated as well as by the most simple means: to give one instance:—in many diœcious plants pollen is carried from the male to the female by the wind, which is perhaps the simplest method conceivable, or by the adherence of the grains to the hairy bodies of insects, which is a method only a little less simple; but in Catasetum the conveyance is effected by the most complex machinery; for in this orchid we have sensitive horns which when touched cause a membrane to rupture, and this sets free certain springs by which the pollen-masses are shot forth like an arrow, and they adhere to the insect's body by a peculiar viscid matter, and then by the breaking of an elastic thread of the right strength the pollen is left sticking to the stigma of the female plant. The complexity of the means used in this and in many other cases, in fact depends on all the previous stages through which the species has passed, and on the successive adaptations of each part during each stage to changed conditions of life.
As some authors consider reciprocal dimorphism to be the first step toward diœciousness, the difficulty of understanding how a trimorphic plant like Lythrum salicaria could become diœcious should be noticed; and as dimorphism and trimorphism are so closely allied, it is not probable that either state is necessarily in any way related to a separation of the sexes—though it may occasionally lead to this end. As far as Lythrum salicaria is concerned, the one tendency which we can discover is towards the abortion of the two sets of stamens in the mid-styled form. This tendency is evinced by its pollen, though abundant and apparently good, yielding a smaller percentage of seed than does the pollen of the corresponding stamens in the other two forms; and this
fact is in itself curious, and shows by what insensibly graduated steps nature moves. If this tendency were carried out the mid-styled form would become a female, depending for its fertilization on two sets of stamens in the long- and short-styled forms; and these two forms would reciprocally fertilize each other like the two forms of Primula or Linum; but there would be no approach to a diœcious condition.
As the case of the trimorphic species of Lythrum is so complicated, and as it is easier to perceive the relations of the sexes in the animal than in the vegetable kingdom, it may be worth while to give, before concluding, a somewhat elaborate simile. We may take the case of a species of Ant, and suppose all the individuals invariably to live in three kinds of communities; in the first, a large-sized female (not to specify other differences) living with six middle-sized and six small-sized males; in the second, a middle-sized female with six large- and six small-sized males; and in the third community, a small-sized female with six large- and six middle-sized males. Each one of these three females, though enabled to unite with any male, would be nearly sterile with her own two sets of males, and likewise with two other sets of males living in the other two communities; for she would be fully fertile only when paired with a male of her own size. Hence the thirty-six males, distributed by half-dozens in the three communities, would be divided into three sets of a dozen each; and these sets, as well as the three females, would differ from each other sexually in exactly the same manner as distinct species of the same genus. Moreover the two sets of males living in the community of the extraordinarily fertile middle-sized female would be less potent sexually than the males of corresponding size in the two other communities. Lastly, we should find that from the eggs laid by each of the three females, all three sorts of females and all three sorts of males were habitually reared—proving to demonstration that all belonged to one and the same species.
To appreciate fully this remarkable case of the reciprocally trimorphic species of Lythrum, we may take a glance at the two great kingdoms of nature and search for anything analogous. With animals we have the most astonishing diversity of structure in the so-called cases of alternate generation, but as such animals have not arrived at maturity, they are not properly comparable with the forms of Lythrum. With mature animals we have extreme differences in structure in the two sexes; we have in
some of the lower animals males, females, and hermaphrodites of the same species; we have the somewhat more curious case of certain Cirripedes which are hermaphrodites, but are sexually aided by whole clusters of what I have called complemental males;1 we have, as Mr. Wallace has lately shown, the females of certain Lepidoptera existing under three distinct forms; but in none of these cases is there any reason to suspect that there is more than one female or one male sexual element. With certain insects, as with Ants, in which there exist, besides males and females, two or three castes of workers, we have a slightly nearer approach to our case, for the workers are so far sexually affected as to have been rendered sterile. With plants, at least with phanerogamic plants, we have not that wonderful series of successive developmental forms so common with animals; nor could this be expected, as plants are fixed to one spot from their birth, and must be adapted throughout life to the same conditions. With plants we have sexual differences in structure, but apparently less strongly marked than with animals, from causes which are in part intelligible, such as there being no sexual selection; again, we have that class of dimorphic flowers so ably discussed recently by Hugo von Mohl, in which some of the flowers are minute, imperfectly developed, and necessarily self-fertile, whilst others are perfect and capable of crossing with other flowers of the same species; but in these several cases we have no reason to suspect that there is more than one female or one male sexual element. When we come to the class of reciprocally dimorphic plants, such as Primula, Linum, &c., we first meet with two masculine and two feminine sexes. But these cases, which seemed only a short time since so strange, now sink almost into insignificance before that of the trimorphic species of Lythrum.
Naturalists are so much accustomed to behold great diversities of structure associated with the two sexes, that they feel no surprise at the fact; but differences in sexual nature have been thought to be the very touchstone of specific distinction. We now see that such sexual differences—the greater or less power of fertilizing and being fertilized—may characterize and keep separate the coexisting individuals of the same species, in the same manner as they characterize and have kept separate those groups of individuals, produced from common parents during the lapse of ages or in different regions, which we rank and denominate as distinct species.
1 Living Cirripedia (1851): p. 55.
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Interested in applying to York as a college/university transfer applicant or a mature student? Then this session is for you!
Join us to learn more about our programs, admission requirements and next steps for transfer and mature applicants. Sessions begin promptly at 6:30 pm.
Please note that this session is for undergraduate programs at York and will not cover information specific to graduate studies.

subset: commoncrawl


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Sources: Illinois Comes to Terms with Groce
The Chicago Tribune and CBSSports.com are reporting that Illinois has agreed to a deal with Ohio University's John Groce to replace Bruce Weber as men's basketball coach.
CBSSports.com, citing an unidentified source, reports that Groce is expected to meet with his team in Athens, Ohio, Thursday and will be introduced at a news conference in Champaign later in the week.
The Tribune, also citing an unidentified source, reports that a disagreement on terms of the contract delayed the hiring for a few days.
Groce has been at Ohio since 2008, leading the Bobcats of the Mid-American Conference to the NCAA tournament twice, including a run to the round of 16 this year that ended with an overtime loss to North Carolina.
Weber was fired after nine seasons at Illinois.
The news means the U of I could be filling vacancies on back-to-back days. Matt Bollant was announced as the new women's basketball coach Wednesday, leaving the University of Wisconsin-Green Bay after a 31 and 2 mark.


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BreadMan wrote:btw, brickshelf is taking forever to moderate my folder, so I put the vids up on youtube:
Brickshelf never makes folders with videos in them public (IIRC). The only videos that they'll make public are .gif videos. I think this is because Kevin Loch does all the moderating himself, and it would take too long for him to watch all the videos.


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Drop the Bass Not The Cash with Bass Drop
Posted by Carter Dotson on May 23rd, 2012
iPhone App - Designed for iPhone, compatible with iPad
Want to make bass drops that are so sick, the CDC will have to declare a pandemic? Want to make dubstep so dirty that you'll feel compelled to put on an episode of Game of Thrones just to feel clean again? Then Bass Drop is a must-download. This app lets anyone make some sick dubstep beats. I'm an undercut away from being Skrillex!
There's a keyboard for making assorted "wub wub wub wub" sounds from 3 different sets, and 12 buttons that can add effects, bass lines, and drum loops. These buttons can be configured with different types of the sounds, and with the ability to have them be toggles, or only play when physically held down. 3 different preference layouts can be saved to make recreating favorite tracks easier. Bass Drop is currently free through the end of May, so aspiring electronic musicians can get in on portable dubstepping at an unbeatable price.
Bass Drop
iPhone App - Designed for iPhone, compatible with iPad
Released: 2012-05-18 :: Category: Game


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At the Detroit auto show, Toyota Division Group Vice President and General Manager Bill Fay announces 100 percent of Corollas sold in America will be built in North America. (Video)
Bill Fay, Toyota group vice president and general manager, says the Corolla Furia Concept features a dramatically swept windshield, sloped roofline and pronounced fender flares. (Video)
Copyright Notice
All materials on this site are for editorial use only. The use of these materials for advertising, marketing or any other commercial purpose is prohibited. They may be cropped but not otherwise modified. To download these materials, you must agree to abide by these terms.


text: This is example home design photos gallery about modern and minimalist house refurbishment design ideas designed by Icon Architects. The new innovation of house designs idea with luxury interior design styles. The refurbishment of this Victorian home is centering at kitchen additional room and garden landscaping. There are great glassy sliding doors at the kitchen as relation to garden. The house garden landscape itself planned as tropical garden planting idea through shiny swimming pool. This outdoor and indoor and outdoor living room transformation makes exclusive atmosphere as well as create the residence sense freshening. At first floor, roof terrace built through seating and BBQ quarter. Below the sample photos of refurbishment house design Innovation ideas that can be your house design inspiration.


text: A Tradition of Academic Excellence Regis University continues to be recognized as a national leader in quality higher education. We are proud of our reputation and our ongoing commitment to encourage each student to explore, dream and discover a lifetime of personal and professional opportunities. Awards and Honors U.S. News & World Report Regis University has been ranked by U.S. News & World Report as a "Top University in the West" for 18 consecutive years. Additionally, Regis University is proud to have achieved the following 2012 "Top Online Education Program Rankings" recognitions: Regis University was included on the "2014 Best Online Graduate Computer Information Technology Programs" list by U.S. News & World Report. Our online bachelor's degree programs ranked in the top four percent for Teaching Practices and Student Engagement and in the top 25 percent for Student Services and Technology Our online master's degree programs in Education ranked in the top 10 percent for Teaching Practices and Student Engagement and in the top 20 percent for Student Services and Technology Our online Master's degree programs in Business ranked in the top 25 percent for Student Services and Technology SuperScholar Smart Choice Ranking The online bachelor's degree programs from Regis University College for Professional Studies were recognized by SuperScholar in its inaugural Smart Choice ranking of the 25 Best Online Colleges and Universities of 2012. Regis University was ranked sixth among the regionally accredited schools with online undergraduate degree programs in the United States. Among the criteria considered were academic quality, student support services, and credibility and prestige in the marketplace. GI Jobs Top Military Friendly School Regis University College for Professional Studies has been named a Top Military Friendly School for 2013 by GI Jobs. U.S. President's Higher Education Community Service Honor Roll In March 2013, Regis University was named to the 2013 U.S. President's Higher Education Community Service Honor Roll with Distinction by the Corporation for National & Community Service. Association of Jesuit Colleges and Universities Conference on Multicultural Affairs In June 2008, Regis University was the recipient of the Distinguished Program/Project Award and the Distinguished Program/Project Award Certificate of Merit by the Association of Jesuit Colleges and Universities Conference on Multicultural Affairs. Peter F. Drucker Foundation Award Presented with the Peter F. Drucker Foundation Award for Innovative Programs. GetEducated.com GetEducated.com's Top 39 Best Buy Online Masters Degrees – Master of Nonprofit Management School of Computer and Information Sciences received a five year (2007–2012) certification from the National Security Agency's National Center of Excellence in Information Assurance. Recognitions Featured in Tom Peters' newsletter "On Achieving Excellence" and on the Wall Street Journal's front page for commitment to quality education for working professionals. Lauded as "One of the best in America at tending to the needs of adult learners" in the classic book What America Does Right by Robert H. Waterman (W. W. Norton & Company) Loretto Heights School of Nursing was recognized by the National Association of Neonatal Nurses (NANN) for "creating an environment that fosters superior leadership skills in neonatal nursing practice."


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Wed, 01 August 2007 at 1:42 pm
Fashion Faceoff: Leona Edmiston Dress
Catherine Bell dresses up in a "Scarlett Style" Leona Edmiston dress for her appearance on FOX's The Morning Show with Mike and Juliet. The Army Wives actress finished off her look with simple red sandals and a tan handbag.
Britney Spears dined in the same floral print dress at Cravings restaurant on Sunset Plaza last week. The pop princess completed her look with fire engine red accessories — head scarf, handbag and heels.
Who wore it best — CATHERINE BELL or BRITNEY SPEARS?
Just Jared on Facebook
Photos: INFphoto.com, Gabo/Dean/Chris/Raul/Bauer-Griffin Posted to: Britney Spears, Catherine Bell, Fashion Faceoff
Sponsored Links by ZergNet


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Alert 2006-49
Subject: Counterfeit Cashier's Checks
Date: August 31, 2006
Description: Rabobank, N.A.
Counterfeit Cashier's Checks of Rabobank, N.A., El Centro, CA
To: Chief Executive Officers of All National Banks; All State Banking Authorities; Chairman, Board of Governors of the Federal Reserve System; Chairman, Federal Deposit Insurance Corporation; Director, Office of Thrift Supervision; Conference of State Bank Supervisors; Deputy Comptrollers (districts); Assistant Deputy Comptrollers; District Counsel; and All Examining Personnel
The above-named bank has reported that counterfeit The above-named bank has reported that counterfeit cashier's checks issued in the bank's name and using a correct routing number of 121139711 are being presented for payment nationwide in connection with a lottery scam.
The volume of these counterfeit items has increased along with the variation of the appearance of the items. Bank officials, therefore, are encouraged to contact Rabobank, N.A. to verify the authenticity of the item that is being presented.
Correspondence accompanying the counterfeit items may contain the letterhead of "American Consumers Sweepstake Lottery" and may be signed by a person purporting to be the president of this organization. The letters notify the recipient of their winning status and instruct them to contact an "agent" at 1-800-963-5144 ext 1 or 647-430-9696 ext 1. The letter further indicates that the internal revenue officer handling the file is Mrs. Carol Francis, who can be reached at 514-827-8898. According to this correspondence, the check is included to pay the IRS non-resident fees.
For additional information and guidance, please contact Patricia Martinez, Rabobank, N.A., 301 Main Street, Salinas, CA 93901 or via telephone at (831) 422-6642.
Additional information concerning this matter that you believe should be brought to the attention of the OCC may be forwarded to:
E-mail: email@example.com
For additional information regarding other types of financial fraud, please visit the OCC's anti-fraud resource at:
Daniel P. Stipano


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Friday, June 18, 2010
Some perspective, please
7 comments:
- Steve Bates said...
Assassination of American citizens without trial, by executive order... DADT still in place... Gitmo still open, with a possibility of moving the whole thing within continental U.S. boundaries... millions of people uninsured AFTER the health care "reform" bill became law... largely ineffective response to the worst environmental disaster in U.S. history... need I go on? and do we have a problem here?
I give Obama credit for one difference from GWB, and it's a big one: Obama is more dependable on women's rights. As an old Planned Parenthood contractor, I appreciate that. But I'm waiting to see a few of those campaign promises, if not implemented, at least acted upon... hell, at least not outright reversed.
Constitutional lawyer, MFA...
- Kath黃rinA俊宇 said...
- This comment has been removed by a blog administrator.
- The Truffle said...
Where did you read about the assassination of citizens? I've been following Glenn Greenwald and don't remember him writing about this.
- The Truffle said...
PS: Check out PoitiFact, which keeps track of Obama's promises.
- Steve Bates said...
Since Greenwald seems to be a source we both trust, please read his post "Presidential assassinations of U.S. citizens." By no means is this episode in January the only instance; I'll leave the finding of the rest to you and Teh Google. Obama obviously feels, as did Bush before him, that the President, acting alone, can make the determination of who is guilty of terrorism and who should die for it, without any sort of due process or trial. I disagree with him in the strongest terms: our system of justice, whether it succeeds or fails in convicting a terrorist, is of far greater significance than the conviction of any particular individual. In the case of an American citizen, there is no constitutional excuse whatsoever for omitting any aspect of due process... none.
- Steve Bates said...
Re: PolitiFact... I just read through a large number of the tracked promises Obama has made (I didn't make it through all the "In the Works" promises; there are simply too many for one individual in one evening) and came to the conclusion that PolitiFact is missing a very salient point: not one promise I read, zip-zero-zilch, had anything to do with restoration of a presidential commitment to civil liberties, civil rights, the propriety of trials, the separation of powers, etc. etc. Such promises are simply not there.
I have to assume that's because Obama wants to keep the powers Bush assumed through sheer executive overreach, the perverse forms of "trial" (e.g., military commissions) devised in the Bush era that practically guarantee conviction of everyone accused of terrorism, and (as noted above) the "right" of the President to order an American citizen abroad assassinated without any form of trial or other due process. I'm sorry, this is just plain wrong, and Obama's pursuit of it is egregious. It outweighs every good thing Obama has done. And this by an alleged constitutional scholar... what an utter embarrassment it is, or ought to be, for him to proceed in this way.
- Jolly Roger said...
No, he certainly isn't a Chimpy, but he doesn't seem to have the respect for due process that I had hoped for.
Honestly.... we either restore the rule of law in this country, or this country will be done. And it'll be sooner, rather than later.


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Beautiful Full Figure Woman: Shar Jackson (Niecy From Moesha) On Celebrity Fit Club! (Shocked To See Her Baby Daddy On The Show Too, Tears Up)
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NEW YORK — The creator of "The Sopranos" said at James Gandolfini's funeral that the actor brought the traits of a sad boy, "amazed and confused," to the role of Tony Soprano.
"You were a good boy," David Chase said Thursday at the ceremony at New York's Cathedral Church of Saint John the Divine.
One of four speakers at the funeral, Chase gave his remarks in the form of a letter to Gandolfini. The actor's widow, Deborah Lin Gandolfini, and two family friends were also speakers at the ceremony.
Chase remembered that Gandolfini once told him that "you know what I want to be? A man. That's all. I want to be a man." Chase said he marveled upon hearing that, since Gandolfini was a man so many others looked up to.
Chase added that, paradoxically, he always felt that in Gandolfini he was seeing a young boy as well as a man.
"A sad boy, amazed and confused," he said. "You could see it in your eyes. That's why you were a great actor."
The 51-year-old actor best known for his role as mob boss Tony Soprano in the HBO series died of a heart attack last week while vacationing with his son in Italy.
Celebrities and fellow actors were among the mourners, along with members of the public who wanted to salute Gandolfini's work.
Those from "The Sopranos" included Edie Falco, Joe Pantoliano, Dominic Chianese, Steve Schirripa, Aida Turturro, Vincent Curatola, Tony Sirico, Lorraine Bracco and Michael Imperioli. Yet another former castmate, Steve Buscemi, chatted with talk show host-comedian Dick Cavett before the ceremony started.
Others from the acting community included Julianna Margulies, Alec Baldwin, Chris Noth, Marcia Gay Harden and Steve Carell. New Jersey Gov. Chris Christie was also on hand.
Some 1,500 seats had been set up in the huge sanctuary. A private family wake was held for the actor Wednesday in New Jersey.
Broadway theaters paid tribute by dimming their lights briefly Wednesday night. Gandolfini was nominated for a Tony Award in 2009 as an actor in "God of Carnage."
Susan Aston, who was Gandolfini's longtime dialogue coach and collaborator, spoke at the funeral of how the actor was devoted to his craft.
"He worked hard," Aston said. "He was disciplined. He studied his roles and did his homework." But when the cameras rolled, his performance would become an act of faith that carried him, she said, "to an uncharted place."
New Jersey accents were easy to hear among members of the public waiting outside the cathedral and waiting for a chance to get in. A few people spoke in Italian.
"I'm a fan," said Saul Stein, 60. "I came to pay my respects today because he's a character I identify with, a family man."
One casual meeting with Gandolfini was enough to bring Robin Eckstein to the funeral.
"I had friends that worked with him," she said. "I had the pleasure of meeting him a few times and he was just lovely. So warm … As soon as he knew you were a friend of a friend you were his friend too. He'll be missed. I missed a meeting at work today. I told them I had a funeral to go to."
Meanwhile, directly across Amsterdam Avenue, in the window of a bar, a large photograph of a grinning Gandolfini was on display, accompanied by a handwritten message that spoke for the actor's fans and friends. It said, simply, "Thank you."
Associated Press correspondent Bethan McKernan and Television Writer David Bauder contributed to this report.


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Last month's employment report showed a gain of 217,000 positions, but job seekers are still likely to find a highly-competitive labor market as hiring prospects remain drab.
Continue Reading Below
But Joanna Weidenmiller puts the blame on the country's hiring system. So she launched a business she claims fixes it.
Weidenmiller, the CEO and co-founder of The One-Page Company, says companies need to find ways to engage job candidates beyond their resumes.
"Resumes only tell the past of a person, which is never enough to predict future performances," she says. "They also can't tell who the candidates really are, and what they are capable of."
The Silicon Valley-based company's 1-Page platform allows employers to have candidates compete for jobs by solving problems in real time that are specific to the job they want.
For example, a company in California has applicants for a sales director position solve the following challenge:
"We are looking to expand into the education sector with force in 2014, and needs to create $2.5 million in sales in 2014. Your challenge is to suggest an idea that reflects your experience and your ability to execute that idea."
The company launched in 2011, and has worked with more than 20 large multi-national corporations with more than 10,000-plus employees, as well as other businesses with between 500 and 5,000 workers, according to Weidenmiller.
"The result is a win-win for employers and candidates," Weidenmiller says. "Companies get unique data they have never had before—predictive data for enhanced decision making on talent. Candidates have the chance to pitch their value beyond their past accomplishments."
Last year, Weindemiller says 1-Page found that 6.8 billion resumes were sent to five million companies, adding that big companies like Google (NASDAQ: GOOG) are getting 70,000 resumes a week. Candidates know their odds are slim, so they apply to as many companies as they can in hopes of increasing their chances of landing an in-person interview.
"To manage this kind of traffic of applications, companies are equipped with resume-processing programs that look for key words and most of the time; they reject resumes only because of the format."
Roy Cohen, career counselor and author of "The Wall Street Professional's Survival Guide," says the process is similar to an "in-basket," an exercise challenge that used to part of the interview process.
"Technology makes this process efficient, and any process that offers additional insight into how a candidate will perform is worth every penny," Cohen says.
He agrees that the online application system doesn't provide a real way for candidates to stand out."Individuals should know it's to their benefit to participate in a challenge like this," Cohen says.
Companies that use 1-Page challenge candidates with real-time problems, and applicants that are really motivated will accept the challenge, Weindemiller says. Candidates can be sourced from LinkedIn (NASDAQ:LKND), internal referrals and more.
"They will do their homework and prove their value with unique ideas and solutions that the companies will never find on a resume," she says. "Those who were not motivated in the first place will soon drop out and focus on the company they really want to work for."
Aside from the skills gap that Weidenmiller says is hurting the job market, she adds understanding and engaging the Millennial workforce is also a big hurdle for employers.
She says her 1-Page platform helps better attract younger workers because they are a "social generation."
"Millennials are soon to be 50% of the workforce in the U.S.," she says. "1-Page engages Millennials by asking their ideas and how they will solve companies' problems, having the chance to design their next job based on what they propose. Suddenly candidates have the power, as they really are those with the value that companies are looking to acquire."


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This is the live worship and sermon broadcast for www.onechurch.tv in Clarksville, Tennessee. Join us at 9am and 11am on Sundays CST. If you have any questions during the service, please contact us by using this link to send an email or by phone/text at (931) 614-0339.
You can also look up the sermon text and submit prayer requests by opening up our event on youversion.com/live


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We know that lots of people are in the unfortunate situation of needing to do a short sale on their home in the Los Angeles area so we are going to do research to find out how to best do a short sale in Los Angeles.
How to do the short sale without being taken advantage of, with reliable agents, who are on your side, who care about you and can understand all that you are going through.
Stay tuned.


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Thank you jb, and you are, I am sure and certain, not alone in those sentiments.
I always sign the liability disclaimer and the terms & conditions form (usually advocating a buddy system) offered to me at the start of any dive trip, but after a six hour session in the witness box I always now add the caveat that "I can in no way be held responsible for the well-being of any other person in the water."
What does that mean? I dive alone - although there may be other people in the water near me..


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Former All Black Sonny Bill Williams admits the chance to win Olympic gold could tempt him to switch to rugby Sevens for the 2016 Rio Games as he weighs up his future in the boxing ring.
The code-hopping New Zealander is about to resume his rugby league career with the Sydney Roosters after winning a 10-round heavyweight boxing bout against veteran South African Frans Botha earlier this month.
But Williams, 27, a member of the World Cup-winning All Blacks team of 2011, told the Sydney Morning Herald that playing Sevens for New Zealand at the Rio Games was an appealing prospect.
"To win a gold medal sounds pretty good," he said.
"As a sportsman, if you don't have those sorts of dreams or aspirations then you may as well just stay in bed."
Williams refused to commit to continuing his boxing career after the draining fight won on points against the vastly more experienced Botha earlier this month in Australia.
"I think I am best at putting on the boots and running out on the field to play rugby union or rugby league," said Williams.
"If I had lost that fight I would have given it away because it is just not worth it, but I overcame some hurdles and the reaction of all the people close to me was good so we will assess it when the time is right."
Williams, on the receiving end of some heavy blows from Botha in the final round of the bout, admitted that boxing was a challenge.
"I am not perfect but I am just trying to keep improving and if everyone is saying I have improved then I will just keep plugging away and maybe have another fight," he said.
"But at this stage I am concentrating on playing rugby league and we will assess that after the season. At this stage, I have had enough of boxing for now."
Earlier this month All Black Liam Messam said he was considering switching to Sevens in order to compete for an Olympic gold medal.
The shortened form of the game becomes a full medal sport in the Brazilian city of Rio de Janeiro in 2016, a year after the next Rugby World Cup in England.


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Now that you are a CSC student, you may find that our educational system is a little different from your home. Many new international students find they have to adjust how they prepare and study for their courses. We have provided the following tips to help you get good grades. Many of our international students are excellent scholars and receive top marks each semester. You can be one of them!
To get good grades, you should:
Faculty, staff and administration at CSC are sincerely interested in your academic success. In addition to numerous student support services at CSC, some of our faculty and staff have offered their advice and suggestions for helping you excel at CSC.
"I find students often avoid approaching their professors when they have a question or concern. The most important advice I could give is for students to contact their instructors when they have concerns or questions about a course or assignment. Often a short conversation or an email can clarify the situation. From a professor's perspective it is important to receive student feedback. It is the only way of knowing if the information is being understood or if instructions for an assignment are well defined."—Ms. Tammi Littrel, CSC instructor
"Your success in college depends on a combination of factors: your behaviors in and out of class, your attitude and commitment to working hard, your awareness of key learning strategies, and your commitment to apply and assess these strategies in your daily life. As with most everything in your life, your motto should be, "I'm responsible for my success!" If you put forth the effort to study effectively, the improved skills will soon become a habit and be just as natural as breathing. The result can be better grades, greater knowledge, and higher self-esteem. These skills will serve you well in your professional and personal life."—Ms. Frances Gonzalez, Tutoring Center
Keys to a successful academic year:
"Welcome to Chadron State College! One of the most valuable relationships you can make in college is with your Academic Advisor. At CSC, you will be first assigned to the "START" Office advisors team. This group of professional staff advisors can help with course schedules, dropping and adding, assisting with major changes, and with financial aid and billing questions. During your second semester you will be assigned to a faculty advisor. This person can be very helpful in guiding you through your academic career at CSC and for giving you good information about your chosen major and career field. If you find you don't easily develop a relationship with your advisor, you can request that a new one be assigned. The other thing to understand is that ANY faculty person can be helpful –not necessarily the one to whom you've been assigned. So, don't hesitate to have conversations with faculty who teach in your chosen degree program or your area of interest. Faculty can be one of the most valuable resources on campus for providing information and guidance about their particular academic discipline. And, if they don't know the answer to your question, they will be able to direct you to someone who does."—Ms. Deena Kennell, Advisor