LECTURE XXIIITAXONOMIC ANOMALIES
LECTURE XXIIITAXONOMIC ANOMALIES
TAXONOMIC ANOMALIES
The theory of descent is founded mainly on comparative studies, which have the advantage of affording a broad base and the convincing effect of concurrent evidence brought together from widely different sources. The theory of mutation on the other hand rests directly upon experimental investigations, and facts concerning the actual descent of one form from another are as yet exceedingly rare. It is always difficult to estimate the validity of conclusions drawn from isolated instances selected from the whole range of contingent phenomena, and this is especially true of the present case. Systematic and physiologic facts seem to indicate the existence of universal laws, and it is not probable that the process of production of new species would be different in the various parts of the animal and vegetable kingdoms. Moreover the principle of unit-characters, the preeminent significance of which has come to be more fully recognized of late, is in full harmony [659] with the theory of sudden mutations. Together these two conceptions go to strengthen the probability of the sudden origin of all specific characters.Experimental researches are limited in their extent, and the number of cases of direct observation of the process of mutation will probably never become large enough to cover the whole field of the theory of descent. Therefore it will always be necessary to show that the similarity between observed and other cases is such as to lift above all doubt the assertion of their resulting from the same causes.Besides the direct comparison of the mutations described in our former lectures, with the analogous cases of the horticultural and natural production of species and varieties at large, another way is open to obtain the required proof. It is the study of the phenomena, designated by Casimir de Candolle by the name of taxonomic anomalies. It is the assertion that characters, which are specific in one case, may be observed to arise as anomalies or as varieties in other instances. If they can be shown to be identical or nearly so in both, it is obviously allowable to assume the same origin for the specific character and for the anomaly. In other terms, the specific marks may be considered as having originated according to the laws [660] that govern the production of anomalies, and we may assume them to lie within reach of our experiments. The experimental treatment of the origin of species may also be looked upon as a method within our grasp.The validity and the significance of these considerations will at once become clear, if we choose a definite example. The broadest and most convincing one appears to me to be afforded by the cohesion of the petals in gamopetalous flowers. According to the current views the families with the petals of their flowers united are regarded as one or two main branches of the whole pedigree of the vegetable kingdom. Eichler and others assume them to constitute one branch, and therefore one large subdivision of the system. Bessey, on the other hand, has shown the probability of a separate origin for those groups which have inferior ovaries. Apart from such divergencies the connation of the petals is universally recognized as one of the most important systematic characters.How may this character have originated? The heath-family or the Ericaceae and their nearest allies are usually considered to be the lowest of the gamopetalous plants. In them the cohesion of the petals is still subject to reversionary exceptions. Such cases of atavism may [661] be observed either as specific marks, or in the way of anomalies.Ledum,MonotropaandPyrola, or the Labrador tea, the Indian pipe and wintergreen are instances of reversionary gamopetalism with free petals. In heaths (Erica Tetralix) and in rhododendrons the same deviation is observed to occur from time to time as an anomaly, and even the commonRhododendron ponticumof our gardens has a variety in which the corolla is more or less split. Sometimes it exhibits five free petals, while at other times only one or two are entirely free, the remaining four being incompletely loosened.Such cases of atavism make it probable that the coherence of the petals has originally arisen by the same method, but by action in the opposite direction. The direct proof of this conclusion is afforded by a curious observation, made by Vilmorin upon the bright and large-flowered garden-poppy,Papaver bracteatum. Like all poppies it has four petals, which are free from one another. In the fields of Messrs. Vilmorin, where it is largely cultivated for its seeds, individuals occur from time to time which are anomalous in this respect. They exhibit a tendency to produce connate petals. Their flowers become monopetalous, and the whole strain is designated by the name ofPapaver[662]bracteatum monopetalum. Henry de Vilmorin had the kindness to send me some of these plants, and they have flowered in my garden during several years. The anomaly is highly variable. Some flowers are quite normal, exhibiting no sign of connation; others are wholly gamopetalous, the four petals being united from their base to the very margin of the cup formed. In consequence of the broadness of the petals however, this cup is so wide as to be very shallow.Intermediate states occur, and not infrequently. Sometimes only two or three petals are united, or the connation does not extend the entire length of the petals. These cases are quite analogous to the imperfect splitting of the corolla of the rhododendron. Giving free rein to our imagination, we may for a moment assume the possibility of a new subdivision of the vegetable kingdom, arising from Vilmorin's poppy and having gamopetalous flowers for its chief character. If the character became fixed, so as to lose its present state of variability, such a group of supposititious gamopetalous plants might be quite analogous to the corresponding real gamopetalous families. Hence there can be no objection to the view, that the heaths have arisen in an analogous manner from their polypetalous ancestors. Other species of [663] the same genus have also been recorded to produce gamopetalous flowers, as for instance,Papaver hybridum, by Hoffmann. Poppies are not the sole example of accidental gamopetaly. Linnaeus observed the same deviation long ago forSaponaria officinalis, and since, it has been seen inClematis Vitalbaby Jaeger, inPeltaria alliaceaby Schimper, inSilene annulataby Boreau and in other instances. No doubt it is not at all of rare occurrence, and the origin of the present gamopetalous families is to be considered as nothing extraordinary. It is, as a matter of fact, remarkable that it has not taken place in more numerous instances, and the mallows show that such opportunities have been available at least more than once.Other instances of taxonomic anomalies are afforded by leaves. Many genera, the species of which mainly bear pinnate or palmate leaves, have stray types with undivided leaves. Among the brambles,Rubus odoratusandR. flexuosusmay be cited, among the aralias,Aralia crassifoliaandA. papyrifera, and among the jasmines, the deliciously scented sambac (Jasminum Sambac). But the most curious instance is that of the telegraph-plant, orDesmodium gyrans, each complete leaf of which consists of a large terminal leaflet and two little lateral ones. These latter keep up, [664] night and day, an irregular jerking movement, which has been compared to the movements of a semaphore.Desmodiumis a papilionaceous plant and closely allied to the genusHedysarum, which has pinnate leaves with numerous pairs of leaflets. Its place in the system leaves no doubt concerning its origin from pinnate-leaved ancestors. At the time of its origination its leaves must have become reduced as to the number of the blades, while the size of the terminal leaflet was correspondingly increased.It might seem difficult to imagine this great change taking place suddenly. However, we are compelled to familiarize ourselves with such hypothetical assumptions. Strange as they may seem to those who are accustomed to the conception of continuous slow improvements, they are nevertheless in complete agreement with what really occurs. Fortunately the direct proof of this assertion can be given, and in a case which is narrowly related, and quite parallel to that of theDesmodium, since it affects a plant of the same family. It is the case of the monophyllous variety of the bastard-acacia orRobinia Pseud-Acacia. In a previous lecture we have seen that it originated suddenly in a French nursery in the year 1855. It can be propagated by seed, and exhibits a curious degree [665] of variability of its leaves. In some instances these are one-bladed, the blade reaching a length of 15 cm., and hardly resembling those of the common bastard-acacia. Other leaves produce one or two small leaflets at the base of the large terminal one, and by this contrivance are seen to be very similar to those of theDesmodium, repeating its chief characters nearly exactly, and only differing somewhat in the relative size of the various parts. Lastly real intermediates are seen between the monophyllous and the pinnate types. As far as I have been able to ascertain, these are produced on weak twigs under unfavorable conditions; the size of the terminal leaflet decreases and the number of the lateral blades increases, showing thereby the presence of the original pinnate type in a latent condition.The sudden origin of this "one-leaved" acacia in a nursery may be taken as a prototype of the ancient origin ofDesmodium. Of course the comparison only relates to a single character, and the movements of the leaflets are not affected by it. But the monophylly, or rather the size of the terminal blade and the reduction of the lateral ones, may be held to be sufficiently illustrated by the bastard-acacia. It is worth while to state, that analogous varieties have also arisen in other genera. The "one-leaved" [666] strawberry has already been referred to. It originated from the ordinary type in Norway and at Paris. The walnut likewise, has its monophyllous variety. It was mentioned for the first time as a cultivated tree about 1864, but its origin is unknown. A similar variety of the walnut, with "one-bladed" leaves but of varying shapes, was found wild in a forest near Dieppe in France some years ago, and appeared to be due to a sudden mutation.Something more is known concerning the "one-bladed" ashes, varieties of which are often seen in our parks and gardens. The common form has broad and deeply serrate leaves, which are far more rounded than the leaflets of the ordinary ash. The majority of the leaves are simple, but some produce one or two smaller leaflets at their base, closely corresponding in this respect to the variations of the "one-bladed" bastard-acacia, and evidently indicating the same latent and atavistic character. In some instances this analogy goes still further, and incompletely pinnate leaves are produced with two or more pairs of leaflets. Besides this variable type another has been described by Willdenow. It has single leaves exclusively, never producing smaller lateral leaflets, and it is said to be absolutely constant from seed, while the more variable types [667] seem to be also more inconstant when propagated sexually. The difference is so striking and affords such a reliable feature that Koch proposed to make two distinct varieties of them, calling the pure typeFraxinus excelsior monophylla, and the varying treesF. excelsior exheterophylla. Some writers, and among them Willdenow, have preferred to separate the "one-leaved" forms from the species, and to call themFraxinus simplicifolia.According to Smith and to Loudon, the "one-leaved" ashes are found wild in different districts in-England. Intermediate forms have not been recorded from these localities. This mode of origin is that already detailed for the laciniate varieties of alders and so many other trees. Hence it may be assumed that the "one-leaved" ashes have sprung suddenly but frequently from the original pinnate species. The pure type of Willdenow should, in this case, be considered as due to a slightly different mutation, perhaps as a pure retrograde variety, while the varying strains may only be eversporting forms. This would likewise explain part of their observed inconstancy.In this respect the historic dates, as collected by Korshinsky, are not very convincing. Vicinism has of course, almost never been excluded, and part of the multiformity of the offspring [668] must obviously be due to this most universal agency. Indirect vicinism also plays some part, and probably affords the explanation of some reputed mutative productions of the variety. So, for instance, in the case of Sinning, who after sowing the seeds of the common ash, got as large a proportion as 2% of monophyllous trees in a culture of some thousand plants. It is probable that his seeds were taken partly from normal plants, and partly from hybrids between the normal and the "one-bladed" type, assuming that these hybrids have pinnate leaves like their specific parent, and bear the characters of the other parent only in a latent condition.Our third example relates to peltate leaves. They have the stalk inserted in the middle of the blade, a contrivance produced by the connation of the two basal lobes. The water-lilies are a well known instance, exhibiting sagittate leaves in the juvenile stage and changing in many species, into nearly circular peltate forms, of whichVictoria regiais a very good example, although its younger stages do not always excite all the interest they deserve. The Indian cress (Tropaeolum), the marsh pennywort orHydrocotyle, and many other instances could be quoted. Sometimes the peltate leaves are not at all orbicular, but are elongated, oblong or elliptic, and with only the lobes [669] at the base united. The lemon-scentedEucalyptus citriodorais one of the most widely known cases. In other instances the peltate leaves become more or less hollow, constituting broad ascidia as in the case of the crassulaceous genusUmbilicus.This connation of the basal lobes is universally considered as a good and normal specific character. Nevertheless it has its manifest analogy in the realm of the anomalies. This is the pitcher or ascidium. On some trees it is of quite common occurrence, as on the lime-tree (Tilia parvifolia) and the magnolia (Magnolia obovataand its hybrids). It is probable that both these forms have varieties with, and others without, ascidia. Of the lime-tree, instances are known of single trees which produce hundreds of such anomalous leaves yearly, and one such a tree is growing in the neighborhood of Amsterdam at Lage Vuursche. I have alluded to these cases more than once, but on this occasion a closer inspection of the structure of the ascidium is required. For this purpose we may take the lime-tree as an example. Take the shape of the normal leaves in the first place. These are cordate at their base and mainly inequilateral, but the general shape varies to a considerable extent. This variation is closely related to the position of the leaves on the twigs, and shows [670] distinct indications of complying with the general law of periodicity. The first leaves are smaller, with more rounded lobes, the subsequent leaves attain a larger size, and their lobes slightly change their forms. In the first leaves the lobes are so broad as to touch one another along a large part of their margins, but in organs formed later this contact gradually diminishes and the typical leaves have the lobes widely separated. Now it is easily understood that the contact or the separation of the lobes must play a part in the construction of the ascidia, as soon as the margins grow together. Leaves which touch one-another, may be affected by the connation without any further malformation. They remain flat, become peltate and exhibit a shape which in some way holds a middle position between the pennyworts and the lemon-scented eucalyptus. Here we have the repetition of the specific characters of these plants by the anomaly of another. Whenever the margins are not in contact, and become connate, notwithstanding their separation, the blade must be folded together in some slight degree, in order to produce the required contact. This is the origin of the ascidium. It is quite superfluous to insist upon the fact that their width or narrowness must depend upon the corresponding normal form. The more distant the [671] lobes, the deeper the ascidium will become. It should be added that this explanation of the different shapes of ascidia is of general validity.Ascidia of the snake-plantain orPlantago lanceolataare narrow tubes, because the leaves are oblong or lanceolate, while those of the broad leaved species of arrowhead, as for instance, theSagittaria japonica, are of a conical shape.From the evidence of the lime-tree we may conclude that normal peltate leaves may have originated in the same way. And from the fact that pitchers are one of the most frequent anomalies, we may conclude that the chance of producing peltate leaves must have been a very great one, and wholly sufficient to account for all observed cases. In every instance the previously existing shape of the leaf must have decided whether peltate or pitcher-like leaves would be formed. As far as we can judge peltate anomalies are quite uninjurious, while ascidia are forms which must impede the effect of the light on the leaf, as they conceal quite an important part of the upper surface. In this way it is easily conceivable that peltate leaves are a frequent specific character, while ascidia are not, as they only appear in the special cases of limited adaptation, as in the instances of the so called pitcher-plants. The generaNepenthes, [672]Sarraceniaand some others are very well known and perhaps even the bladderworts orUtriculariamight be included here.The reproduction of specific characters by anomalous ascidia is not at all limited to the general case as described above. More minute details may be seen to be duplicated in the same way. Proofs are afforded on one side by incomplete ascidia, and on the other by the double cups.Incomplete ascidia are those of theNepenthes. The leaf is divided into three parts, a blade, a tendril and the pitcher. Or in other words, the limb produces a tendril at its summit, by means of which the plant is enabled to fasten itself to surrounding shrubs and to climb between their branches. But the end of this tendril bears a well-formed urn, which however, is produced only after the revolving and grasping movements of the tendril have been made. Some species have more rounded and some more elongated ascidia and often the shape is seen to change with the development of the stem. The mouth of the urn is strengthened by a thick rim and covered with a lid. Numerous curious contrivances in these structures to catch ants and other insects have been described, but as they have no relation to our present discussion, we shall abstain from dealing with them. [673] Likewise we must refrain from a consideration of the physiologic qualities of the tendril, and confine our attention to the combination of a limb, a naked midvein and an ascidium. This combination is to be the basis of our discussion. It is liable to be produced all of a sudden. This assertion is proved by its occurrence as a varietal mark in one of our most ordinary cultivated plants. It is the group known asCroton, belonging to the genusCodiaeum. A variety is calledinterruptumand anotherappendiculatum, and these names both relate to the interruption of the leaves by a naked midvein. The leaves are seen to be built up of three parts. The lower half retains the aspect of a limb; it is crowned by a vein without lateral nerves or blade-like expansions, and this stalk in its turn bears a short limb on its summit. The base of this apical limb exhibits two connate lobes, forming together a wide cup or ascidium. It should be stated that theseinterruptumvarieties are highly variable, especially in the relative size of the three principal parts of the leaf. Though it is of course conceded that the ascidium ofNepentheshas many secondary devices which are lacking inCroton, it seems hardly allowable to deny the possibility of an analogous origin for both. Those of theCroton, according to our knowledge regarding similar cases, must [674] have arisen at once, and hence the conclusion that the ascidia ofNepenthesare also originally due to a sudden mutation. Interrupted leaves, with an ascidium on a naked prolongation of the midvein, are by no means limited to theCrotonvarieties. As stray anomalies they have often been observed, and I myself had the opportunity of collecting them on magnolia, on clover and on some other species. They are additional evidence in support of the explanation given above.In the same way double ascidia may be made use of to explain the foliar cups of the teasels and some other plants, as for instance, some European snakeroots (Eryngium maritimumandE. campestre), or the floral leaves of the honeysuckle. The leaves on the stems of the teasels are disposed in pairs, and the bases of the two leaves of each pair are connate so as to constitute large cups. We have already mentioned these cups, and recall them in the present connection to use them as a prototype of the double ascidia. These are constituted of two opposite leaves, accidentally connated at their base or along some part of their margins. If the leaves are sessile, the analogy with the teasels is complete, as shown, for instance, in a case ofCotyledon, a crassulaceous plant which is [675] known to produce such cups from time to time. They are narrower than those of the teasel, but this depends, as we have seen for the "one-leaved" ascidium, on the shape of the original leaf. In other respects they exactly imitate the teasel cups showing thereby how these cups may probably have originated.In numerous cases of anomalies some accidental structures are parallel to specific characters, while others are not, being obviously injurious to their bearers. So it is also with the double ascidia. In the case of stalked leaves the two opposite stalks must, of course, constitute a long and very narrow tube, when growing together. This tube must bear at its summit the conical ascidium produced by the two connate limbs. At its base however, it includes the terminal bud of the stem, and frequently the tube is so narrow as to impede its further development. By this contrivance the double ascidium assumes a terminal position. Instances have been observed on magnolia, inBoehmeriaand in other cases.Flowers on leaves are of rare occurrence. Notwithstanding this, they constitute specific characters in some instances, accidental anomalies in others.Helwingia rusciflora florais the most curious and best known instance. It is a little shrub, belonging to the Cornaceae, and [676] has broad elliptical undivided leaves. On the middle of the midvein these leaves are seen to bear small clusters of flowers; indeed this is the only place where flowers are produced. Each cluster has from 13-15 flowers, of which some are staminate and borne on stalks, while others are pistillate and nearly sessile. These flowers are small and of a pale greenish color and yield small stone-fruits, with a thin coating of pulpy tissue. As the name indicates, this mode of flowering is closely similar to that ofRuscus, which however, does not bear its flowers and berries on real leaves, but on leaflike expansions of the twigs.Phyllonoma ruscifolia, a saxifragaceous plant, bears the same specific name, indicating a similar origin of the flowers. Other instances have been collected by Casimir de Candolle, but their number is very small.As a varietal mark, flowers on leaves likewise rarely occur. One instance however, is very remarkable, and we have already dealt with it, when treating of constant varieties, and of the lack of vicinism in the case of species with exclusive self-fertilization.It is the "Nepaul-barley" orHordeum trifurcatum. The leaves, which in this case bear the adventitious flowers, are the inner scales of the spikelets, and not on green leaves as in the [677] cases already alluded to. But this of course makes no real difference. The character is variable to a high degree, and this fact indicates its varietal nature, though it should be recalled that at least with theHelwingia, the majority of the leaves are destitute of flowers, and that in this way some degree of variability is present in this normal case too.All in all there are three sorts of "Nepaul-barley." They have the same varietal mark, but belong to different species of barley. These are differentiated according to the number of the rows in which the grains are seen on the spikes. These numbers may be two, four or six, giving rise to the specific names ofHordeum distichum,tetrastichumandhexastichum. Whether these three varieties are of independent, but parallel origin, or are to be considered as due to a single mutation and subsequent crosses is not known, all of them being of ancient origin. Historic evidence concerning their birth is wholly wanting. From analogy it would seem probable that the character had arisen by a mutation in one of the three named species, and had been transferred to others by means of accidental crosses, even as it has been artificially transmitted of late to quite a number of other sorts. But however admissible this conception may seem, there is of course no real objection [678] to the assumption of independent and parallel mutations.For the purpose of a comparison with theHelwingiatype we are however, not at all concerned with the species to which thetrifurcatumvariety belongs, but only with the varietal mark itself. The spikelets may be one-, two- or three-flowered, according to the species. If we choose for further consideration thehexastichumtype, each spikelet produces three normal flowers and afterwards three normal grains. Morphologically however, the spikelet is not homologous to those parts of other grasses which have the same name. It is constituted of three real spikelets, and thus deserves the name of a triple construction. Each of these three little organs has its normal pair of outer scales or glumae. These are linear and short, ending in a long and narrow spine. Those of the middle-most spikelets stand on its outer side, while those of the lateral part are placed transversely. In this way they form a kind of involucre around the central parts. The latter consist of the inner and outer palets or scales, each two of which include one of the flowers. The outer palet is to be considered as the metamorphosed leaf, in the aril of which the flower is produced. In the common sorts of barley it bears a long awn, giving thereby its typical aspect to the [679] whole spike. The axillary flower is protected on the opposite side by a two-keeled inner palet. Each flower exhibits three stamens and an ovary. In the six-rowed barley all the three flowers of a triple spikelet are fertile, and each of them has a long awn on the top of the outer palet. But in the two-rowed species only the middle-most flower is normal and has an awn, the two remaining being sterile and more or less rudimentary and with only very short awns. From this description it is easily seen that the species of barley may be distinguished from one another, even at a casual glance, by the number of the rows of the awns, and therefore by the shape of the entire spikes. This striking feature, however, does not exist in the "Nepaul-barley." The awns are replaced by curiously shaped appendices, which are three-lobed. The central lobe is oblong and hollow, and forms a kind of hood, which covers a small supernumerary floret. The two lateral lobes are narrower, often linear and extended into a smaller or longer awn. These awns are mostly turned away from the center of the spike. The central lobe may sometimes bear two small florets, but ordinarily only one is to be found, and this is often incomplete, having only one or two stamens, or is different in some other way. [680] These narrow lateral lobes heighten the abnormal aspect of the whole spike.They are only produced at a somewhat advanced stage of the development of the palet, are united to one another and to the central part by strong veins, which form transversal anastomoses at their insertion. The length of these awns is very variable, and this quality is perhaps the most striking of the whole variety. Often they reach only 1-2 mm., or the majority may become longer and attain even 1 cm., while here and there, between them, longer ones are inserted, extending in some instances even as far as 3 cm. from the spike. Their transverse position in such cases is strikingly contrasted with the ordinary erect type of the awns.These lateral lobes are to be regarded, from the morphologic point of view, as differentiated parts of the blade of the leaf. Before they are formed, or coincidently with the beginning of their development, the summit of the central lobe becomes hollow, and the development of the supernumerary flower commences. In different varieties, and especially in the most recent crosses of them, this development is excessively variable.The accidental flower arises at some distance beneath the summit of the scale, on its middle [681] vein. The development begins with the protrusion of a little scale, and the flower itself is situated beneath this scale, and is to be protected by it and by the primary scale, but is turned upside down at the same time. Opposite to this organ, which represents the outer palet of the adventitious flower, two little swollen bodies are evolved. In the normal flowers of barley and other grains and grasses their function is to open the flowers by swelling, and afterwards collapse and allow them to close.In the adventitious flowers of the "Nepaul-barley," however, this function is quite superfluous. The stamens occur in varying numbers; typically there are three, but not rarely less, or more, are seen. In some instances the complete double whorl of six, corresponding to the ancestral monocotyledonous type, has been found. This is a very curious case of systematic atavism, quite analogous to theIris pallida abavia, previously alluded to, which likewise has six stamens, and to the cases given in a previous lecture. But for our present discussion it is of no further interest. The ovary is situated in the middle of the flower, and in some instances two have been observed. This is also to be considered as a case of atavism.All these parts of the adventitious flower are more or less subject to arrest of development, [682] in a later stage. They may even sometimes become abnormal. Stamens may unite into pairs, or carpels bear four stigmas. The pollen-sacs are as a rule barren, the mother-cells undergoing atrophy, while normal grains are seen but rarely. Likewise the ovaries are rudimentary, but Wittmack has observed the occasional production of ripe grains from these abnormal florets.The scale is seldom seen to extend any farther upwards than the supernumerary flower. But in the rare instances where it does prolong its growth, it may repeat the abnormality and bear a second floret above the first. This of course is generally much weaker, and more rudimentary.Raciborsky, who has lately given a full and very accurate description of this anomaly, lays great stress upon the fact that it is quite useless. It is perhaps the most obviously useless structure in the whole vegetable kingdom. Notwithstanding this, it has come to be as completely hereditary as any of the most beautiful adaptations in nature. Therefore it is one of the most serious objections to the hypothesis of slow and gradual improvements on the sole ground of their usefulness. The struggle for life and natural selection are manifestly inadequate to give even the slightest indication of [683] an explanation of this case. It is simply impossible to imagine the causes that might have produced such a character. The only way out of this difficulty is to assume that it has arisen at once, in its present apparently differentiated and very variable condition, and that, being quite uninjurious and since it does not decrease the fertility of the race, it has never been subjected to natural selection, and so has saved itself from destruction.But if we once grant the probability of the origin of the "Nepaul-barley" by a sudden mutation, we obviously must assume the same in the case of theHelwingiaand other normal instances. In this way we gain a further support for our assertion, that even the strangest specific characters may have arisen suddenly.After having detailed at some length those proofs which seem to be the most striking, and which had not been previously described with sufficient detail, we may now take a hasty survey of other contingent cases. In the first place the cruciate flowers of some onagraceous plants should be remembered. Small linear petals occur as a specific character inOenothera cruciataof the Adirondacks, but have been seen to arise as sudden mutations in the common evening-primrose (O. biennis) in Holland, and in the willow-herb (Epilobium hirsutum) in England. [684] Leaves placed in whorls of three are very rare. The oleander, juniper and some few other plants have ternate whorls as a specific character. As an anomaly, ternate whorls are far more common, and perhaps any plant with opposite leaves may from time to time produce them. Races rich in this abnormality are found in the wild state in the yellow loosestrife orLysimachia vulgaris, in which it is a very variable specific character, the whorls varying from two to four leaves. In the cultivated state it is met with in the myrtle orMyrtus communis, where it has come to be of some importance in Israelitic ritual. Crisped leaves are known in a mallow,Malva crispa, and as a variety in cabbages, parsley, lettuce and others. The orbicular fruits of Heeger's shepherd's purse (Capsella heegeri) recall similar fruits of other cruciferous genera, as for instance,Camelina. Screw-like stems with wide spirals are specific in the flower-stalks ofCyclamenandVallisneria, varietal inJuncus effusus spiralisand accidental inScirpus lacustris. Dormant buds or small bulbs in inflorescences are normal for wild onions,Polygonum viviparumand others, varietal inPoa alpina viviparaand perhaps inAgave vivipara, and accidental in plantains (Plantago lanceolata),Saxifraga umbrosaand others. [685] Cleft leaves, one of the most general anomalies, are typical inBoehmeria biloba. The adnation of the peduncles of the inflorescences to the stem is typical inSolanumand accidental in many other cases.It seems quite superfluous to add further proof. It is a very general phenomenon that specific characters occur in other genera as anomalies, and under such circumstances that the idea of a slow evolution on the ground of utility is absolutely excluded. No other explanation remains than that of a sudden mutation, and once granted for the abnormal cases, this explanation must obviously likewise be granted for the analogous specific characters.Our whole discussion shows that mutations, once observed in definite instances, afford the most probable basis for the explanation of specific characters at large.
[686]
LECTURE XXIVTHE HYPOTHESIS OF PERIODIC MUTATIONS
LECTURE XXIVTHE HYPOTHESIS OF PERIODIC MUTATIONS
THE HYPOTHESIS OF PERIODIC MUTATIONS
The prevailing belief that slow and gradual, nearly invisible changes constitute the process of evolution in the animal and vegetable kingdom, did not offer a strong stimulus for experimental research. No appreciable response to any external agency was of course to be expected. Responses were supposed to be produced, but the corresponding outward changes would be too small to betray themselves to the investigator.The direct observation of the mutations of the evening-primrose has changed the whole aspect of the problem at once. It is no longer a matter dealing with purely hypothetical conditions. Instead of the vague notions, uncertain hopes, and a priori conceptions, that have hitherto confused the investigator, methods of observation have been formulated, suitable for the attainment of definite results, the general nature of which is already known.To my mind the real value of the discovery [687] of the mutability of the evening-primrose lies in its usefulness as a guide for further work. The view that it might be an isolated case, lying outside of the usual procedure of nature, can hardly be sustained. On such a supposition it would be far too rare to be disclosed by the investigation of a small number of plants from a limited area. Its appearance within the limited field of inquiry of a single man would have been almost a miracle.The assumption seems justified that analogous cases will be met with, perhaps even in larger numbers, when similar methods of observation are used in the investigation of plants of other regions. The mutable condition may not be predicated of the evening-primroses alone. It must be a universal phenomenon, although affecting a small proportion of the inhabitants of any region at one time: perhaps not more than one in a hundred species, or perhaps not more than one in a thousand, or even fewer may be expected to exhibit it. The exact proportion is immaterial, because the number of mutable instances among the many thousands of species in existence must be far too large for all of them to be submitted to close scrutiny.It is evident from the above discussion that next in importance to the discovery of the prototype of mutation is the formulation of methods [688] for bringing additional instances to light. These methods may direct effort toward two different modes of investigation. We may search for mutable plants in nature, or we may hope to induce species to become mutable by artificial methods. The first promises to yield results most quickly, but the scope of the second is much greater and it may yield results of far more importance. Indeed, if it should once become possible to bring plants to mutate at our will and perhaps even in arbitrarily chosen directions, there is no limit to the power we may finally hope to gain over nature.What is to guide us in this new line of work? Is it the minute inspection of the features of the process in the case of the evening-primroses? Or are we to base our hopes and our methods on broader conceptions of nature's laws? Is it the systematic study of species and varieties, and the biologic inquiry into their real hereditary units? Or is the theory of descent to be our starting-point? Are we to rest our conceptions on the experience of the breeder, or is perhaps the geologic pedigree of all organic life to open to us better prospects of success?The answer to all such questions is a very simple one. All possibilities must be considered, and no line of investigation ignored. For myself I have based my field-researches and my [689] testing of native plants on the hypothesis of unit-characters as deduced from Darwin'sPangenesis. This conception led to the expectation of two different kinds of variability, one slow and one sudden. The sudden ones known at the time were considered as sports, and seemed limited to retrograde changes, or to cases of minor importance. The idea that sudden steps might be taken as the principal method of evolution could be derived from the hypothesis of unit characters, but the evidence might be too remote for a starting point for experimental investigation.The success of my test has given proof to the contrary. Hence the assertion that no evidence is to be considered as inadequate for the purpose under discussion. Sometime a method of discovering, or of producing, mutable plants may be found, but until this is done, all facts of whatever nature or direction must be made use of. A very slight indication may change forever the whole aspect of the problem.The probabilities are now greatly in favor of our finding out the causes of evolution by a close scrutiny of what really happens in nature. A persistent study of the physiologic factors of this evolution is the chief condition of success. To this study field-observations may contribute as well as direct experiments, [690] microscopical investigations as well as extended pedigree-cultures. The cooperation of many workers is required to cover the field. Somewhere no doubt the desired principle lies hidden, but until it is discovered, all methods must be tried.With this conception as the best starting point for further investigation, we may now make a brief survey of the other phase of the problem. We shall try to connect our observations on the evening-primroses with the theory of descent at large.We start with two main facts. One is the mutability of Lamarck's primrose, and the second is the immutable condition of quite a number of other species. Among them are some of its near allies, the common and the small flowered evening-primrose, orOenothera biennisandO. muricata.From these facts, a very important question arises in connection with the theory of descent. Is the mutability of our evening-primroses temporary, or is it a permanent condition? A discussion of this problem will give us the means of reaching a definite idea as to the scope of our inquiries.Let us consider the present first. If mutability is a permanent condition, it has of course no beginning, and moreover is not due to the [691] agency of external circumstances. Should this be granted for the evening-primrose, it would have to be predicated for other species found in a mutable state. Then, of course, it would be useless to investigate the causes of mutability at large, and we should have to limit ourselves to the testing of large numbers of plants in order to ascertain which are mutable and which not.If, on the other hand, mutability is not a permanent feature, it must once have had a beginning, and this beginning itself must have had an external cause. The amount of mutability and its possible directions may be assumed to be due to internal causes. The determination of the moment at which they will become active can never be the result of internal causes. It must be assigned to some external factor, and as soon as this is discovered the way for experimental investigation is open.In the second place we must consider the past. On the supposition of permanency all the ancestors of the evening-primrose must have been mutable. By the alternative view mutability must have been a periodic phenomenon, producing at times new qualities, and at other times leaving the plants unchanged during long successions of generations. The present mutable state must then have been preceded by an immutable [692] condition, but of course thousands of mutations must have been required to produce the evening-primroses from their most remote ancestors.If we take the species into consideration that are not mutable at present, we may ask how we are to harmonize them with each of the two theories proposed. If mutability is permanent, it is manifest that the whole pedigree of the animal and vegetable kingdom is to be considered as built up of main mutable lines, and that the thousands of constant species can only be taken to represent lateral branches of the genealogic tree.These lateral branches would have lost the capacity of mutating, possessed by all their ancestors. And as the principle of the hypothesis under discussion does not allow a resumption of this habit, they would be doomed to eternal constancy until they finally die out. Loss of mutability, under this conception, means loss of the capacity for all further development. Only those lines of the main pedigree which have retained this capacity would have a future; all others would die out without any chance of progression.If, on the other hand, mutability is not permanent, but a periodic condition, all lines of the genealogic tree must be assumed to show alternatively [693] mutating and constant species. Some lines may be mutating at the present moment; others may momentarily be constant. The mutating lines will probably sooner or later revert to the inactive state, while the powers of development now dormant may then become awakened on other branches.The view of permanency represents life as being surrounded with unavoidable death, the principle of periodicity, on the contrary, follows the idea of resurrection, granting the possibility of future progression for all living beings. At the same time it yields a more hopeful prospect for experimental inquiry.Experience must decide between the two main theories. It demonstrates the existence of polymorphous genera, such asDrabaandViolaand hundreds of others. They clearly indicate a previous state of mutability. Their systematic relation is exactly what would be expected, if they were the result of such a period. Perhaps mutability has not wholly ceased in them, but, might be found to survive in some of their members. Such very rich genera however, are not the rule, but are exceptional cases, indicating the rarity of powerful mutative changes.On the other hand, species may remain in a state of constancy during long, apparently during indefinite, ages.[694] Many facts plead in favor of the constancy of species. This principle has always been recognized by systematists. Temporarily the current form of the theory of natural selection has assumed species to be inconstant, ever changing and continuously being improved and adapted to the requirements of the life-conditions. The followers of the theory of descent believed that this conclusion was unavoidable, and were induced to deny the manifest fact that species are constant entities. The mutation theory gives a clew to the final combination of the two contending ideas. Reducing the changeability of the species to distinct and probably short periods, it at once explains how the stability of species perfectly agrees with the principle of descent through modification.On the other hand, the hypothesis of mutative periods is by no means irreconcilable with the observed facts of constancy. Such casual changes can be proved by observations such as those upon the evening-primrose, but it is obvious that a disproof can never be given. The principle grants the present constancy of the vast majority of living forms, and only claims the exceptional occurrence of definite changes.Proofs of the constancy of species have been given in different ways. The high degree of similarity of the individuals of most of our [695] species has never been denied. It is observed throughout extended localities, and during long series of years. Other proofs are afforded by those plants which have been transported to distant localities some time since, but do not exhibit any change as a result of this migration. Widely dispersed plants remain the same throughout their range, provided that they belong to a single elementary species. Many species have been introduced from America into Europe and have spread rapidly and widely. The Canadian horsetail (Erigeron canadensis), the evening-primrose and many other instances could be given. They have not developed any special European features after their introduction. Though exposed to other environmental conditions and to competition with other species, they have not succeeded in developing a new character. Such species as proved adequate to the new environment have succeeded, while those which did not have succumbed.Much farther back is the separation of the species which now live both in arctic regions and on the summits of our highest mountaintops. If we compare the alpine flora with the arctic plants, a high degree of similarity at once strikes us. Some forms are quite identical; others are slightly different, manifestly representing elementary species of the same systematic [696] type. Still others are more distant or even belong to different genera. The latter, and even the diverging, though nearly allied, elementary species, do not yield adequate evidence in any direction.They may as well have lived together in the long ages before the separation of the now widely distant floras, or have sprung from a common ancestor living at that time, and subsequently have changed their habits. After excluding these unreliable instances, a good number of species remain, which are quite the same in the arctic and alpine regions and on the summits of distant mountain ranges. As no transportation over such large distances can have brought them from one locality to the other, no other explanation is left than that they have been wholly constant and unchanged ever since the glacial period which separated them. Obviously they must have been subjected to widely changing conditions. The fact of their stability through all these outward changes is the best proof that the ordinary external conditions do not necessarily have an influence on specific evolution. They may have such a result in some instances, in others they obviously have not. Many arctic forms bearing the specific name ofalpinusjustify this conclusion.Astragalus alpinus,Phleum alpinum,Hieracium alpinumand [697] others from the northern parts of Norway may be cited as examples.Thus Primula imperialis has been found in the Himalayas, and many other plants of the high mountains of Java, Ceylon and northern India are identical forms. Some species from the Cameroons and from Abyssinia have been found on the mountains of Madagascar. Some peculiar Australian types are represented on the summit of Kini Balu in Borneo. None of these species, of course, are found in the intervening lowlands, and the only possible explanation of their identity is the conception of a common post-glacial origin, coupled with complete stability. This stability is all the more remarkable as nearly allied but slightly divergent forms have also been reported from almost all of these localities. Other evidence is obtained by the comparison of ancient plants with their living representatives. The remains in tombs of ancient Egypt have always afforded strong support of the views of the adherents of the theory of stability, and to my mind they still do so. The cereals and fruits and even the flowers and leaves in the funeral wreaths of Rameses and Amen-Hotep are the same that are still now cultivated in Egypt. Nearly a hundred or more species have been identified. Flowers ofAcacia, leaves ofMimusops, [698] petals ofNymphaeamay be cited as instances, and they are as perfectly preserved as the best herbarium-specimens of the present time. The petals and stamens retain their original colors, displaying them as brightly as is consistent with their dry state.Paleontologic evidence points to the same conclusion. Of course the remains are incomplete, and rarely adequate for a close comparison. The range of fluctuating variability should be examined first, but the test of elementary species given by their constancy from seed cannot, of course, be applied. Apart from these difficulties, paleontologists agree in recognizing the very great age of large numbers of species. It would require a too close survey of geologic facts to go into details on this point. Suffice it to say that in more recent Tertiary deposits many species have been identified with living forms. In the Miocene period especially, the similarity of the types of phanerogamic plants with their present offspring, becomes so striking that in a large number of cases specific distinctions rest in greater part on theoretical conceptions rather than on real facts. For a long time the idea prevailed that the same species could not have existed through more than one geologic period. Many distinctions founded on this belief have since had to be abandoned. [699] Species of algae belonging to the well-preserved group of the diatoms, are said to have remained unchanged from the Carboniferous period up to the present time.Summing up the results of this very hasty survey, we may assert that species remain unchanged for indefinite periods, while at times they are in the alternative condition. Then at once they produce new forms often in large numbers, giving rise to swarms of subspecies. All facts point to the conclusion that these periods of stability and mutability alternate more or less regularly with one another. Of course a direct proof of this view cannot, as yet, be given, but this conclusion is forced upon us by a consideration of known facts bearing on the principle of constancy and evolution.If we are right in this general conception, we may ask further, what is to be the exact place of our group of new evening-primroses in this theory? In order to give an adequate answer, we must consider the whole range of the observations from a broader point of view. First of all it is evident that the real mutating period must be assumed to be much longer than the time covered by my observations. Neither the beginning nor the end have been seen. It is quite obvious thatOenothera lamarckianawas in a mutating condition when I first [700] saw it, seventeen years ago. How long had it been so? Had it commenced to mutate after its introduction into Europe, some time ago, or was it already previously in this state? It is as yet impossible to decide this point. Perhaps the mutable state is very old, and dates from the time of the first importation of the species into Europe.Apart from all such considerations the period of the direct observations, and the possible duration of the mutability through even more than a century, would constitute only a moment, if compared with the whole geologic time. Starting from this conception the pedigree of our mutations must be considered as only one small group. Instead of figuring a fan of mutants for each year, we must condense all the succeeding swarms into one single fan, as might be done also forDraba vernaand other polymorphous species. InOenotherathe main stem is prolonged upwards beyond the fan; in the others the main stem is lacking or at least undiscernable, but this feature manifestly is only of secondary importance. We might even prefer the image of a fan, adjusted laterally to a stem, which itself is not interrupted by this branch.On this principle two further considerations are to be discussed. First the structure of the [701] fan itself, and secondly the combination of succeeding fans into a common genealogic tree.The composition of the fan as a whole includes more than is directly indicated by the facts concerning the birth of new species. They arise in considerable quantities, and each of them in large numbers of individuals, either in the same or in succeeding years. This multiple origin must obviously have the effect of strengthening the new types, and of heightening their chances in the struggle for life. Arising in a single specimen they would have little chance of success, since in the field among thousands of seeds perhaps one only survives and attains complete development. Thousands or at least hundreds of mutated seeds are thus required to produce one mutated individual, and then, how small are its chances of surviving! The mutations proceed in all directions, as I have pointed out in a former lecture. Some are useful, others might become so if the circumstances were accidentally changed in definite directions, or if a migration from the original locality might take place. Many others are without any real worth, or even injurious. Harmless or even slightly useless ones have been seen to maintain themselves in the field during the seventeen years of my research, as proved byOenothera laevifoliaandOenothera[702]brevistylis. Most of the others quickly disappear.This failure of a large part of the productions of nature deserves to be considered at some length. It may be elevated to a principle, and may be made use of to explain many difficult points of the theory of descent. If, in order to secure one good novelty, nature must produce ten or twenty or perhaps more bad ones at the same time, the possibility of improvements coming by pure chance must be granted at once. All hypotheses concerning the direct causes of adaptation at once become superfluous, and the great principle enunciated by Darwin once more reigns supreme.In this way too, the mutation-period of the evening-primrose is to be considered as a prototype. Assuming it as such provisionally, it may aid us in arranging the facts of descent so as to allow of a deeper insight and a closer scrutiny. All swarms of elementary species are the remains of far larger initial groups. All species containing only a few subspecies may be supposed to have thrown off at the outset far more numerous lateral branches, out of which however, the greater part have been lost, being unfit for the surrounding conditions. It is the principle of the struggle for life between elementary species, followed by the survival of the [703] fittest, the law of the selection of species, which we have already laid stress upon more than once.Our second consideration is also based upon the frequent repetition of the several mutations. Obviously a common cause must prevail. The faculty of producingnanellaorlataremains the same through all the years. This faculty must be one and the same for all the hundreds of mutative productions of the same form. When and how did it originate? At the outset it must have been produced in a latent condition, and even yet it must be assumed to be continuously present in this state, and only to become active at distant intervals. But it is manifest that the original production of the characters ofOenothera gigaswas a phenomenon of far greater importance than the subsequent accidental transition of this quality into the active state. Hence the conclusion that at the beginning of each series of analogous mutations there must have been one greater and more intrinsic mutation, which opened the possibility to all its successors. This was the origination of the new character itself, and it is easily seen that this incipient change is to be considered as the real one. All others are only its visible expressions.Considering the mutative period of our evening-primrose [704] as one unit-stride section in the great genealogic tree, this period includes two nearly related, but not identical changes. One is the production of new specific characters in the latent condition, and the other is the bringing of them to light and putting them into active existence. These two main factors are consequently to be assumed in all hypothetic conceptions of previous mutative periods.Are all mutations to be considered as limited to such periods? Of course not. Stray mutations may occur as well. Our knowledge concerning this point is inadequate for any definite statement. Swarms of variable species are easily recognized, if the remnants are not too few. But if only one or two new species have survived, how can we tell whether they have originated-alone or together with others. This difficulty is still more pronounced in regard to paleontologic facts, as the remains of geologic swarms are often found, but the absence of numerous mutations can hardly be proved in any case.I have more than once found occasion to lay stress on the importance of a distinction between progressive and retrograde mutations in previous lectures. All improvement is, of course, by the first of these modes of evolution, but apparent losses of organs or qualities are [705] perhaps of still more universal occurrence. Progression and regression are seen to go hand in hand everywhere. No large group and probably even no genus or large species has been evolved without the joint agency of these two great principles. In the mutation-period of the evening-primroses the observed facts give direct support to this conclusion, since some of the new species proved, on closer inspection, to be retrograde varieties, while others manifestly owe their origin to progressive steps. Such steps may be small and in a wrong direction; notwithstanding this they may be due to the acquisition of a wholly new character and therefore belong to the process of progression at large.Between them however, there is a definite contrast, which possibly is in intimate connection with the question of periodic and stray mutations. Obviously each progressive change is dependent upon the production of a new character, for whenever this is lacking, no such mutation is possible. Retrograde changes, on the other hand, do not require such elaborate preliminary work. Each character may be converted into the latent condition, and for all we know, a special preparation for this purpose is not at all necessary. It is readily granted that such special preparation may occur, because the [706] great numbers in which our dwarf variety of theOenotheraare yearly produced are suggestive of such a condition. On the other hand, thelaevifoliaandbrevistylismutations have not been repeated, at least not in a visible way.From this discussion we may infer that it is quite possible that a large part of the progressive changes, and a smaller part of the retrograde mutations, are combined into groups, owing their origin to common external agencies. The periods in which such groups occur would constitute the mutative periods. Besides them the majority of the retrograde changes and some progressive steps might occur separately, each being due to some special cause. Degressive mutations, or those which arise by the return of latent qualities to activity, would of course belong with the latter group.This assumption of a stray and isolated production of varieties is to a large degree supported by experience in horticulture. Here there are no real swarms of mutations. Sudden leaps in variability are not rare, but then they are due to hybridization. Apart from this mixture of characters, varieties as a rule appear separately, often with intervals of dozens of years, and without the least suggestion of a common cause. It is quite superfluous to go into details, as we have dealt with the horticultural [707] mutations at sufficient length on a previous occasion. Only the instance of the peloric toadflax might be recalled here, because the historic and geographic evidence, combined with the results of our pedigree-experiment, plainly show that peloric mutations are quite independent of any periodic condition. They may occur anywhere in the wide range of the toad-flax, and the capacity of repeatedly producing them has lasted some centuries at least, and is perhaps even as old as the species itself.Leaving aside such stray mutations, we may now consider the probable constitution of the great lines of the genealogic tree of the evening primroses, and of the whole vegetable and animal kingdom at large. The idea of drawing up a pedigree for the chief groups of living organisms is originally due to Haeckel, who used this graphic method to support the Darwinian theory of descent. Of course, Haeckel's genealogic trees are of a purely hypothetic nature, and have no other purpose than to convey a clear conception of the notion of descent, and of the great lines of evolution at large. Obviously all details are subject to doubt, and many have accordingly been changed by his successors. These changes may be considered as partial improvements, and the somewhat picturesque form of Haeckel's pedigree might well be replaced by [708] more simple plans. But the changes have by no means removed the doubts, nor have they been able to supplant the general impression of distinct groups, united by broad lines. This feature is very essential, and it is easily seen to correspond with the conception of swarms, as we have deduced it from the study of the lesser groups.Genealogic trees are the result of comparative studies; they are far removed from the results of experimental inquiry concerning the origin of species. What are the links which bind them together? Obviously they must be sought in the mutative periods, which have immediately preceded the present one. In the case of the evening-primrose the systematic arrangement of the allied species readily guides us in the delimitations of such periods. For manifestly the species of the large genus ofOenotheraare grouped in swarms, the youngest or most recent of which we have under observation. Its immediate predecessor must have been the subgenusOnagra, which is considered by some authors as consisting of a single systematic species,Oenothera biennis. Its multifarious forms point to a common origin, not only morphologically but also historically. Following this line backward or downward we reach another apparent mutation-period, which includes the origin of [709] the group calledOenothera, with a large number of species of the same general type as theOnagra-forms, Still farther downward comes the old genusOenotheraitself, with numerous subgenera diverging in sundry characters and directions.Proceeding still farther we might easily construct a main stem with numerous succeeding fans of lateral branches, and thus reach, from our new empirical point of view, the theoretical conclusion already formulated.Paleontologic facts readily agree with this conception. The swarms of species and varieties are found to succeed one another like so many stories. The same images are repeated, and the single stories seem to be connected by the main stems, which in each tier produce the whole number of allied forms. Only a few prevailing lines are prolonged through numerous geologic periods; the vast majority of the lateral branches are limited each to its own storey. It is simply the extension of the pedigree of the evening-primroses backward through ages, with the same construction and the same leading features. There can be no doubt that we are quite justified in assuming that evolution has followed the same general laws through the whole duration of life on earth. Only a moment of their lifetime is disclosed to us, but it [710] is quite sufficient to enable us to discern the laws and to conjecture the outlines of the whole scheme of evolution.A grave objection which has, often, and from the very outset, been urged against Darwin's conception of very slow and nearly imperceptible changes, is the enormously long time required. If evolution does not proceed any faster than what we can see at present, and if the process must be assumed to have gone on in the same slow manner always, thousands of millions of years would have been needed to develop the higher types of animals and plants from their earliest ancestors.Now it is not at all probable that the duration of life on earth includes such an incredibly long time. Quite on the contrary the lifetime of the earth seems to be limited to a few millions of years. The researches of Lord Kelvin and other eminent physicists seem to leave no doubt on this point. Of course all estimates of this kind are only vague and approximate, but for our present purposes they may be considered as sufficiently exact.In a paper published in 1862 Sir William Thomson (now Lord Kelvin) first endeavored to show that great limitation had to be put upon the enormous demand for time made by Lyell, Darwin and other biologists. From a consideration [711] of the secular cooling of the earth, as deduced from the increasing temperature in deep mines, he concluded that the entire age of the earth must have been more than twenty and less than forty millions of years, and probably much nearer twenty than forty. His views have been much criticised by other physicists, but in the main they have gained an ever-increasing support in the way of evidence. New mines of greater depth have been bored, and their temperatures have proved that the figures of Lord Kelvin are strikingly near the truth. George Darwin has calculated that the separation of the moon from the earth must have taken place some fifty-six millions of years ago. Geikie has estimated the existence of the solid crust of the earth at the most as a hundred million years. The first appearance of the crust must soon have been succeeded by the formation of the seas, and a long time does not seem to have been required to cool the seas to such a degree that life became possible. It is very probable that life originally commenced in the great seas, and that the forms which are now usually included in the plankton or floating-life included the very first living beings. According to Brooks, life must have existed in this floating condition during long primeval epochs, and evolved nearly all the main branches of the animal and vegetable kingdom [712] before sinking to the bottom of the sea, and later producing the vast number of diverse forms which now adorn the sea and land.All these evolutions, however, must have been very rapid, especially at the beginning, and together cannot have taken more time than the figures given above.The agency of the larger streams, and the deposits which they bring into the seas, afford further evidence. The amount of dissolved salts, especially of sodium chloride, has been made the subject of a calculation by Joly, and the amount of lime has been estimated by Eugene Dubois. Joly found fifty-five and Dubois thirty-six millions of years as the probable duration of the age of the rivers, and both figures correspond to the above dates as closely as might be expected from the discussion of evidence so very incomplete and limited.All in all it seems evident that the duration of life does not comply with the demands of the conception of very slow and continuous evolution. Now it is easily seen, that the idea of successive mutations is quite independent of this difficulty. Even assuming that some thousands of characters must have been acquired in order to produce the higher animals and plants of the present time, no valid objection is raised. The demands of the biologists and the results of [713] the physicists are harmonized on the ground of the theory of mutation.The steps may be surmised to have never been essentially larger than in the mutations now going on under our eyes, and some thousands of them may be estimated as sufficient to account for the entire organization of the higher forms. Granting between twenty and forty millions of years since the beginning of life, the intervals between two successive mutations may have been centuries and even thousands of years. As yet there has been no objection cited against this assumption, and hence we see that the lack of harmony between the demands of biologists and the results of the physicists disappears in the light of the theory of mutation.Summing up the results of this discussion, we may justifiably assert that the conclusions derived from the observations and experiments made with evening-primroses and other plants in the main agree satisfactorily with the inferences drawn from paleontologic, geologic and systematic evidence. Obviously these experiments are wonderfully supported by the whole of our knowledge concerning evolution. For this reason the laws discovered in the experimental garden may be considered of great importance, and they may guide us in our further inquiries. Without doubt many minor [714] points are in need of correction and elaboration, but such improvements of our knowledge will gradually increase our means of discovering new instances and, new proofs.The conception of mutation periods producing swarms of species from time to time, among which only a few have a chance of survival, promises to become the basis for speculative pedigree-diagrams, as well as for experimental investigations.
[715]