Saturday, 27 May 2017

Spinach-iron data transformations: Boussingault (1872) to Berg (1913)

For an introduction see any other entry under the spinach-iron label of this blog

In 1872, Jean-Baptiste Boussingault ("Du fer contenu dans le sang et dans les aliments." Comptes Rendus de l'Académie des Sciences, Tome 74: 1353-1359) published his results on the iron contents in the blood of various animals and in food products. This publication contains a table at pp. 1355-56 listing the contents of "Fer exprimé à l'état métallique. Dans 100 grammes de matiére." This means that the values are not for iron oxide (Fe2O3), which was the usual state in which iron contents were measured but elemental. The second statement ("Dans 100 grammes de matiére.") was specified in the text above the table: "En ce qui concerne les aliments, les dosages ont éte exécutés à l'état où ils sont consommés, c'est-à-dire avec leur eau constitutionnelle." (In the case of food, the dosages were carried out in the state in which they were consumed, that is to say with their constitutional water.) Hence, Boussingault's table gives the iron content for spinach leaves ("Feuilles d'épinards") as 0.0045g per 100 gram fresh matter.

In 1897, Emil Häusermann ("Die Assimilation des Eisens. Zeitschrift für Physiologische Chemie 23: 555-592) published a table (pages 586-588) listing the iron contents of food products per 100g dry matter. He cited Boussingault for various items, but the values vary.

Boussingault (g per 100g fresh matter)     Häusermann (mg per 100g dry matter)
Riz: 0.0015 Reis: 1.7
Haricots blancs: 0.0074 Weisse Bohnen: 8.3
Lentilles: 0.0083 Linsen: 9.5
Pommes de terre: 0.0016 Kartoffeln: 6.4
Feuilles d'epinards: 0.0045 Spinat: 39.1

Obviously, Häusermann took values for the water contents of the food stuffs from elsewhere, in order to calculate the iron contents in the dry matter from Boussingault's values for fresh matter. Apparently, these values of the water contents of the fresh matter were 11.7% in rice, 10.8% in white beans, 12.6% in lentil, 75.0% in potatoes and 88.5% in spinach leaves. Bunge (1892. "Weitere Untersuchungen über die Aufnahme des Eisens in den Organismus des Säuglings." Zeitschrift für Physiologische Chemie 16:173-186) has done the same with Boussingault's data before and given König (1889. "Chemie der menschlichen Nahrungsmittel.") as his source for water contents (see footnote ****, here). As Bunge took the water content for spinach to be 88.49%, we can conclude that König has also been Häusermann's source.

In a later edition, Josef König (1904. Chemie der menschlichen Nahrungs- und Genussmittel, Band 2, p. 353) cited the value of Häusermann properly, that is, he gave the range of values as 32.7-39.1mg per 100gr dry matter (the first value being from Bunge 1892 and the second from Häusermann 1897).

Ragnar Berg (1913. Die Nahrungs- und Genussmittel. p. 34-35) gave the contents of iron-oxide (Fe2O3) in 100g fresh matter as he explained in the introduction at page 6: "Damit nun jeder leicht umrechnen kann [...], habe ich in den folgenden Tabellen [...] den Gehalt von 100 g frischen Nahrungsmitteln an einzelnen Mineralbestandteilen in Grammen [...] aufgeführt."
     Berg (1913, p. 34-35) cited König (1904) with a value of 0.0596g Fe2O3 in 100g fresh spinach. (Berg indicated the sources by superscripts given above the values in the table. As he explained at page 11 of the introduction, the roman numeral I stands for König 1904.) If we assume that Berg took the average value (35.9mg/100g dry matter) of König's range (32.7-39.1mg/100g dry matter), then König's average iron (Fe) content in dry matter amounts to 60% of the iron-oxide (Fe2O3) content that Berg imputed to König for fresh matter.
    Berg can hardly have assumed that 60% of fresh spinach leaves were dry matter, when Häusermann had earlier taken its water content to amount to 88.5% (see above). Berg's transformation factor lies much closer to the 70% that is suggested as the correction factor needed to calculate the portion of the mass of Fe2O3 that is due to the iron in it according to the atomic weights (Fe: 55.8; O: 16). Nevertheless, a discrepancy of 10% remains. That is still not satisfying to see how Berg got from König's range of iron contents for dry matter to his imputation to König of iron-oxide content for fresh matter.

Anyway, Berg also performed his own analysis and that yielded 0.0437g Fe2O3 per 100g fresh matter. As it happens, this value was just about ten times higher than what Boussingault had started with (0.0045g Fe per 100g fresh matter) despite the fact that no decimal separator had been misplaced in any of the various data transformations.

Carl von Noorden & Hugo Salomon (1920. Handbuch der Ernährungslehre. Erster Band, p. 476) gave a range of 44-60mg Fe2O3 per 100g spinach and cited Berg (1913) as well as Hermann Schall & August Heisler (1917. Nahrungsmitteltabelle. 5. Auflage. Curt Kabitzsch Verlag, Würzburg) as sources.

The publication of Schall & Heisler (1917, not online) in turn has two values for spinach at page 41: 60mg and 44mg per 100g fresh matter. The latter value bears a footnote referring that value to "R. Berg" (sic), who had given 0.0437g/100g fresh mater as the result of his own analysis. As Berg also gave 0.0596g/100g fresh weight and referred that to König (1904), I presme that Schall & Heisler have simply taken that value from Berg as well, rounded it, but did not specifically cite König, because they have gotten it from a secondary source. The introduction of Schall & Heisler 81917) states that they collected data from "König, Rubner, Atwater und Byrant, Schwenkenbecher, Sautier, Strauss, Tischler, Leva, v. Noorden, Nauny, Magnus-Levy, Janney, Walker Hall, Brugsch, Bessau und Schmidt, Hesse, Offer und Rosenquist, Vogel, Berg, Albu-Neuberg, das "Deutsche Bäderbuch", die Angaben der Nahrungsmittelindustrie u. a. mehr." Sic! No sources, journals, publishers, years or anything else to ease retrival. The publication contains no reference list either to look up citations.

Saturday, 29 April 2017

Comparing Matthew's and Darwin's theories

The following analyses and compares the evolutionary schemes of Patrick Matthew and Charles Darwin.

1. Matthew’s propositions
In his book On Naval Timber and Arboriculture, Patrick Matthew explicitly excluded the wider range of circumstances under cultivation as the cause of increased differences in varieties and attributed it instead to man’s interference with natural selection:
Man’s interference, by preventing this natural process of selection among plants, independent of the wider range of circumstances to which he introduces them, has increased the difference in varieties particularly in the more domesticated kinds.1 (Matthew 1831, 308)
Furthermore, he regarded adaptation to be relative in comparison to competitors as illustrated by examples that ecologists would today call cases of competitive exclusion:
The natural soil and climate of a tree, is often very far from being the soil and climate most suited to its growth, and is only the situation where it has greater power of occupancy, than any other plant whose germ is present. The pines do not cover the pine barrens of America, because they prefer such soil, or grow most luxuriant in such soil; they would thrive much better, that is, grow faster, in the natural allotment of the oak and the walnut, and also mature to a better wood in this deeper richer soil. But the oak and the walnut banish them to inferior soil from greater power of occupancy in good soil, as the pines, in their turn, banish other plants from inferior sands —some to still more sterile location, by the same means of greater powers of occupancy in these sands.”2 (Matthew 1831, 302f, his emphases)
While the oak did better than the pine on rich soils, it fared worse on poor soils. They excluded each other on the respective soils. Without competition, the pine would also do better on rich soils. While evolutionists could now call this idea relative fitness or adaptedness and ecologists could call it reciprocal competitive exclusion (pines from rich soils oaks from poor soils), Matthew called it the power of occupancy.
    However, Matthew did not state this observation of relative fitness, in order to develop an evolutionary argument. On the contrary, the problem at hand was the practical question where to plant different tree species in order to get the best timber. In particular, Matthew (1831, 302f) used his observation to refute Steuart’s claim that trees should always be planted in the soils and climates that resemble their natural habitat. Instead, he maintained that pines would yield better timber if planted in soils that were richer than their natural provenience because they were only excluded from these soils by competitors.
    And further down he used the same logic of this argument, in order to confirm Cruickshank’s claim that the oak could grow throughout Scotland although it does not tend to do so naturally:
The highest latitude to which a tree, or any other kind of plant, reproducing by seed, naturally extends, depending on the ripening of the seed, and also on the power of occupancy, is however different from that where it will grow, when ripe seeds are produced from the coldest place where they ripen, and all the competitors removed; and under the system of shelter belts, hardy pine nurses, and seeds from the nearest place where they ripen, we have no doubt that oaks may be extended to a colder situation than Nature herself would have placed them. (Matthew 1831, 356f)
Again, the problem at hand was not evolutionary, but the practical question whether the area for growing the valuable oak timber could be expanded by a system of getting seeds from the coldest places, where they still ripen, putting them into nursery belts sheltered by hardy pines and removing all competitors from the immediate vicinity.3
    Finally, he explicated his evolutionary ideas in an addendum to the appendix and integrated his concept of relative fitness (power of occupancy) into it as follows:
1. Species are fixed for long periods of time:
A particular conformity, each after its own kind, when in a state of nature, termed species, no doubt exists to a considerable degree. This conformity [of species] has existed during the last forty centuries. (Matthew 1831, 381)
2. Geologists discover like conformities of fossil species within each epoch, but also almost complete difference between the fossil species of one epoch from those of all others (Matthew 1831, 381).
3. This discontinuity (punctuation in the fossil record) is initiated by catastrophes.
The destructive liquid currents […] destroying nearly all living things, must have reduced existence so much, that an unoccupied field would be formed for new diverging ramifications of life […], these remnants, in the course of time, moulding and accommodating their being anew to the change of circumstances, and to every possible means of subsistence, and the millions of ages of regularity which appear to have followed between the epochs, probably after this accommodation was completed, affording fossil deposits of regular specific character. (Matthew 1831, 382f)
4. Pondering the question whether the transformation of species after such a catastrophe was due to repeated new creation, admixture of species, or adaptation (“gradual circumstance-suited modification”), Matthew (1831, 384) opts for the latter.
5. He then, partly, attributes this adaptation to natural selection in a paragraph in which the power of occupancy re-occurs:
The self-regulating adaptive disposition of organized life may, in part, be traced to the extreme fecundity of Nature, who, as before stated, has, in all the varieties of her offspring, a prolific power much beyond (in many cases a thousandfold) what is necessary to fill up the vacancies caused by senile decay. As the field of existence is limited and pre-occupied, it is only the hardier, more robust, better suited to circumstance individuals, who are able to struggle forward to maturity, these inhabiting only the situations to which they have superior adaptation and greater power of occupancy than any other kind; the weaker, less circumstance-suited, being prematurely destroyed. This principle is in constant action, […] in such immense waste of primary and youthful life, those only come forward to maturity from the strict ordeal by which Nature tests their adaptation to her standard of perfection and fitness to continue their kind by reproduction. (Matthew 1831, 384f, my highlighting)
6. Finally, he states how this explains both the conformity (fixity) of species in the periods of stasis as well as their transformation after catastrophic changes:
From the unremitting operation of this law […], a considerable uniformity of figure, colour, and character, is induced, constituting species; the breed gradually acquiring the very best possible [not perfect] adaptation of these to its condition which it is susceptible of, and when alterations of circumstance occur, thus changing in character to suit these as far as its nature is susceptible of change. (Matthew 1831, 385)
The remainder of this addendum is made up of considerations of additional causes of adaptation (hence he traced adaptation only “in part” to natural selection) as well as the anomaly of the human species that either modifies other species to its needs or exterminates them.

2. Darwin’s theory development
Eiseley (1959) interpreted Darwin’s statements from 1844 and 1868 as if they were statements of one and the same theory. Since then, Darwin studies have traced Darwin’s long theory development in minute detail (e.g., Ruse 1975; Browne 1980; Kohn 1980; 1986; 2009; Schweber 1980; Beddall 1988; Mayr 1992; Richardson 1981; Ospovat 1995; Hector and Hooper 2002; Costa 2009; Mannouris 2011; Pearce 2010; Richards 2012). Consequently, Darwin’s earlier statements cannot be taken to be informed by the same theory as his later ones. Before getting into a detailed comparison of Darwin with Matthew, let me briefly sketch Darwin’s intellectual development in a conveniently simplified way. Between his return from the Beagle voyage and 1844, Darwin developed what will be referred to as his early theory of species transformation through natural selection. Between that time and 1856, he was obsessively occupied with taxonomic work concerning small-scale studies of barnacle variation (LIT) as well as large-scale patterns in botanical systematics (Browne 1980). Thereafter, he began to write down his mature theory, posthumously published as Natural Selection (Stauffer 1975), but this was famously interrupted by a letter from from Alfred Russel Wallace precipitating the events that lead to the publication of the Origin of Species as an abstract of the big Natural Selection book he had planned.

2.1 Darwin’s early theory
Despite embracing the idea of species transformation by 1837, he continued to work by theories that agreed with doctrines of natural theology up to the mid-1850s. For example, Darwin (1909[1844]) thought that species hardly varied under natural conditions, because intermarriage would swamp individual variants. He thought that large scale changes in conditions needed to push organisms out of their natural state first. This would affect their reproductive system to increase fortuitous variation in the offspring. Only after this boost of undirected variation could selection adapt the species to the new conditions. Under normal conditions, natural selection was stabilising (keeping species fixed) as generally believed to be true in doctrines of natural theology. Natural selection could only transform species under exceptional conditions.

2.2 Darwin’s later theory
In the mid-1850s Darwin removed the special conditions of existence from the center stage of his theory and allowed that even slight individual differences could be accumulated by natural selection without being swamped. The keystone in this development was his principle of divergence.4 This did not simply explain the long known pattern of diverging ramifications as a contingent result of variation and selection, speciation and extinction, but took divergence as adaptive in itself.5 The sources motivating Darwin’s search for this principle have been traced to studies on embryology, morphology, taxonomy and first ecological experiments by von Baer, Owen, Milne-Edwards and Sinclair respectively (Browne 1980; Kohn 1981; 1986; 2009; Schweber 1980; Beddall 1988; Mayr 1992; Ospovat 1995, chap. 5-8; Hector and Hooper 2002; Pearce 2010; Richards 2012).
    Darwin tried to explain a large-scale pattern of taxonomy that the species within larger genera were also more divergent morphologically than the species in smaller genera. He had invested a lot of research into developing a statistical measure for that pattern and the principle of divergence was his explanation of it (Browne 1980). Starting from an analogy with economic specialisation, Darwin apparently arrived at the conclusion that specialisation is also advantageous for species (Tammone 1995) and construed his principle of divergence as an explanation for the taxonomic pattern from it.
  Darwin (1859, 114, 116, 118, 121), took species to be imperfectly adapted and divergent forms to be improved (better specialised) and therefore to almost necessarily exterminate the intermediate forms, which were usually the own parental forms.
As each species tends by its geometrical ratio of reproduction to increase inordinately in number; and as the modified descendants of each species will be enabled to increase by so much the more as they become more diversified in habits and structure, so as to be enabled to seize on many and widely different places in the economy of nature, there will be a constant tendency in natural selection to preserve the most divergent offspring of any one species. Hence during a long-continued course of modification, the slight differences, characteristic of varieties of the same species, tend to be augmented into the greater differences characteristic of species of the same genus. New and improved varieties will inevitably supplant and exterminate the older, less improved and intermediate varieties; and thus species are rendered to a large extent defined and distinct objects. (Darwin 1859, 470)
The pattern of diverging ramifications was no longer contingent but a necessary result of adaptative change, first, and competitive extinction of intermediate forms thereafter. This idea is no longer valid (Mayr 1992, 354f). Why should selection inevitable preserve the diverging (more specialised) form and exterminate the parental (more generalist) form? The point, here, is not whether Darwin’s principle of divergence is still valid today, but that its development, causal structure and consequences differs from Matthew’s ideas as follows. 
    To distinguish Darwin's early from Darwin's late theory, one could say that the drive for adaptive change was environmental in his early theory, but competitive in his later theory.

3. Theory comparison
Historians of science often claim that Darwin's early theory or theories of others from before The Origin of Species in 1859 included a conception of perfect adaptedness (e.g., Ospovat 1995; Fleming 2013). To address that claim, the meaning of perfect and relative adaptation needs to be clarified first. Perfect adaptation requires two things: (1) A perfectly adapted species will not do better elsewhere and (2) it can also not be ousted from its niche, neither by other species nor by divergent descendants of its own. Suppose a species that was perfectly adapted for one niche but its actual range beyond this spot was restricted by competitors. That would fulfill conditions 1 and 2 from above. But it cannot be Matthew's conception, because such a species would not do better elsewhere.
    Let's stick with Matthew’s example of pines and oaks. If, for example, pines were perfectly adapted for poor soils and cold climate, they would do best there. Rich soils and warmer climate would have adverse physiological effects (similar to over-manuring or desiccation). Pines could still occur on rich soils in the absence of superior competitors, but they would not do as good there as on poor soils. Matthew, however, thins that pines would really do better on rich soils, if only the competitors would not exclude them. Matthew's concept of imperfect adaptation breaks condition 1 for perfection, but Darwin later theory breaks both condition 1 and 2. 
    This explains why Darwin, in his early theory, needed environmental change to alter the conditions to which a species was perfectly adapted and thus induce it to vary markedly, as if under domestication. First the environmental factors change and render a species ill adapted, then it varies markedly, then natural selection works to transform it. Geographic isolation is needed in this early theory, in order to get more than one species.
   The relative adaptation of pines and oaks in Matthew’s scheme also explains why he did not merely need environmental change but catastrophes. The catastrophe needs to exterminate the competitors and leave an unoccupied field of existence into which the pines could then spread and thereafter adapt to the new conditions which they meet there. That is, Matthew's scheme would even work if, no environmental change occurred, but the competitors were nevertheless removed somehow. That’s why he advised to remove the competitors if one wanted to grow oaks in Scotland or pines on rich soil. Conversely, if the soil of the pine habitats became rich and the climate mild, somehow, but that of the oaks would stay as it was, the latter would simply exterminate the former. Hence, catastrophes.
   Imagine a miraculous catastrophe that would exterminate the majority of all species, but leave the environmental conditions on earth just as they were. Such a catastrophe would have the same effect in Matthew's scheme as one that altered the environmental conditions along with exterminating competitors. With perfect adaptation, however, the pines would keep sitting in their niche of poor soils and cold climates and their spread to adjacent habitats would be an extremely slow process. The variant occurring could move only tiny bits beyond the conditions perfect for their ancestors. It would not possibly yield the ultra-rapid adaptive change that Matthew needs in his scheme.6
   In comparison, Matthew's scheme is one of stasis punctuated by catastrophes; in the periods of stasis natural selection keeps the species to their niches. This agrees with the doctrine of natural theology in that natural selection has a species fixing effect during stasis. It disagrees with natural theology in that species are not perfectly adapted to their niche, but only kept there by competitors. Therefore, catastrophes have a species transforming effect by removing competitors. Matthew does not need repeated creation, because the remnant species will naturally spread into the empty fields of existence after a catastrophe. The same natural selection that kept the species fixed now transforms them.
    In Darwin's mature theory, after the principle of divergence, the environmental conditions included other species, but the fiercest competitor of a parental species was the divergent daughter species. That is, the oak was not exterminated by a catastrophe and then the pine moved in, but the parental oak was exterminated by divergent daughter species (other oaks) that were more specialised and therefore better adapted.
    In Matthew's scheme relative adaptedness means that the pine could do better on rich soils and would spread there, if the competitors (hardwood) were removed. In Darwin's later scheme relative adaptedness means that a pine species which could do better on rich soils could also be better adapted for poor soils than it actually is. Therefore divergent descendant species (that are more specialised for poor soils) will oust the parental species. That is how the principle of divergence is the inverse of Matthew's idea of relative power of occupancy. Where Darwin saw sharp wedges packed close together and driven inwards by incessant blows, Matthew saw pointed arrows lashing outwards by incessant throws.
   As the conception of perfect and relative adaptedness is useless for describing Matthew's scheme of evolutionary change, it should be replaced by a conception that distinguishes between an environmental and a competitive drive for adaptive change. We then see that Matthew's theory conceived competition as a stabilising factor and catastrophes as an environmental drive for change by clearing the field from competitors. In Darwin's early conception the environmental change does not drive evolutionary change by removing competitors, but by removing the species from its zone of comfort. 
   In Darwin's late theory, however, competition has become the drive for evolution and environmental change can increase this drive. But competition is never a hindrance to adaptive change in any of Darwin's theories as it is in Matthew's.
    For redundancy, Matthew's concept of relative adaptedness means that the pine is not the species that is best adapted to rich soils and therefore ousted from them, and the oak is not the species best adapted to poor soils and therefore ousted from them. However, although both are only better adapted to their natural place than the competitors—not perfectly adapted—Matthew did not use that, in order to develop an intrinsic evolutionary drive from it (like the principle of divergence). Instead he took on board the external driver of catastrophes.
    For Darwin, the pine is better adapted to poor soils than the oak, okay, but it could also be still better adapted to its natural habitation on poor soils. And Darwin uses that for developing an intrinsic evolutionary drive that needs no recourse to external drives. He used to need the external drive of changed conditions in his early theory, in order to push a species from its comfort zone. But his later theory, while it can accommodate such external factors of evolution, has in in-build drive for adaptive change (the principle of divergence).

4. Conclusion
In conclusion, Matthew’s theory of evolution is a chimera pairing the old doctrine that natural selection (usually) keeps the species fixed with the catastrophism that allowed for rapid transformation and radiation of species by natural selection after a catastrophe. Darwin’s principle of divergence, however, was his final break away from the doctrine that natural selection kept species fixed.
    The principle of divergence differs from Matthew’s ideas in its development (from Darwin’s own studies of systematics and an analogy with economics), its causal structure and its consequences. This constitutes a negative paternity test for Matthew. It exculpates Darwin from claims of plagiarising Matthew for the time after he shifted from his early (1844) to his mature theory (1856-58). His early theory, without the principle of divergence, was equally different from Matthew’s ideas:
Darwin: environmental change renders a species ill adapted → this leads to increased variation → this allows species transformation by natural selection;
Matthew: adaptation is always relative → competition keeps species fixed in their natural place → catastrophe removes competition → species spreads and natural selection transforms it to new conditions. 
This exculpates Darwin for the time from his return from the Beagle voyage till 1844. Hence, the time window left for this particular plagiarism claim is hardly open anymore. 

--- Notes ---
 
1 Note that “difference in varieties” does not necessarily mean individual variability.
2 Prideaux John Selby (1842, 391) cites this passage of Matthew only to refuse it: “Matthew, however, in his able treatise on naval timber seems to think that its indigenous location in such districts arises not so much from preference of soils of the nature above-mentioned, as from its having more power of occupancy in such soils than any other plant of the country; and this opinion he endeavours to support by stating that the Pinus sylvestris, planted in a good or rich soil, attains larger dimensions and its best timber properties, and that it is only driven from this superior soil by the greater power of occupancy, and that it is only driven from this superior soil by the greater power of occupancy possessed by the oak and other deciduous trees, an opinion in which we cannot altogether acquiesce, as we see no reason why the fir, if it grows with such additional vigour in a richer soil, as Mr. Matthew asserts, should, at the same time, be unable to maintain a contest with the oak or other tree.” Apparently, Selby did not comprehend the dynamics of competition and simply assumed that equal size at maturity of pine and oak meant equal competitive powers.
3 Again, he reiterated this logic of relative fitness in his book Emigration Fields Matthew (1839, 38): “The soil of Upper Canada is in many places of fair quality, and generally superior to that of the eastern provinces. The timber consists mostly of large-leafed deciduous trees, while that of the eastern provinces is chiefly pine. This distribution is partly due to soil as well as climate, and has lead to the belief that pines prefer inferior soils. This, however, is not the case. Hard-wood trees have only greater power of occupancy in warmer climates and richer soils and pines in the cold and inferior, especially in the more siliceous.” (see also p. 57)
4 Davies (2013, 727) suggested that Wallace (1856, 214)could have inspired Darwin’s principle of divergence: “It is probable that in very few cases is there a direct affinity between two groups, each being more or less distantly related to some common extinct group, so that we should represent their connexion more accurately by making our central line a blank, for the extinct portion of the group, and placing our families right and left, at different distances from it.”
5 Though anachronistic, the modern distinction between stabilising, directional, and disruptive selection can help understand the principle of divergence. Where, beforehand, the usual circumstances of a species rendered natural selection stabilising, or absent according to Darwin’s early theory, and changed conditions of life were required to render it directional (transforming), disruptive selection for divergence could now also occur under the usual conditions of existence. Transformation, speciation, and extinction were no longer conditional upon large-scale changes of physical circumstances.
6 Matthew (1831) believed that “millions of ages” (p. 383) of stasis, the last one lasting “forty centuries” (p. 381) had been punctuated by an equal number of catastrophes followed by short periods of adaptation and diversification. To cut him some slack, let the average period of stasis be 3000 rather than 4000 years. This will amount to 3 billion years of stasis. Life on earth exists for 3.9 billion years, minus the stasis leaves 900 million years for adaptation. Slotting them in after the catastrophes and before the periods of stasis leaves 900 years, on average, for each period of evolutionary change. And in that time the new fauna and flora had to re-evolve from catastrophes “destroying nearly all living things” (p. 382f). This extremely compressive limitation shows that Matthew’s scheme lacked time for evolutionary change.

--- References ---
  
Beddall, Barbara G. (1988). Darwin and divergence: the Wallace connection. Journal of the History of Biology, 21, 1-68.
Browne, Janet (1980). Darwin’s botanical arithmetic and the principle of divergence, 1854-1858. Journal of the History of Biology, 13, 53-89.
Costa, James T. (2009). The Darwinian revelation: tracing the origin and evolution of an idea. BioScience, 59, 886-894.
Darwin, Charles R. (1859). On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. 1st edition. John Murray, London.
Darwin, Charles R. (1868). The Variation of Animals and Plants under Domestication, Vol. 2. London: John Murray.
Darwin, Charles R. (1909). The Foundations of The Origin of Species. Two Essays written in 1842 and 1844. Edited with an introduction by Francis Darwin. Cambridge University Press.
Davies, Roy (2013). 1 July 1858: what Wallace knew, what Lyell thought he knew; what both he and Hooker took on trust; and what Charles Darwin never told them. Biological Journal of the Linnean Society, 109, 725-736.
Eiseley, Loren C. (1959). Charles Darwin, Edward Blyth, and the Theory of Natural Selection. With an appendix reprinting a Memoir of Edward Blyth by Arthur Grote. Proceedings of the American Philosophical Society, 103(1), 94-158.
Fleming, Leonore (2013) The notion of limited perfect adaptednessin Darwin's principle of divergence. Perspectives on Science 21: 1-22.
Hector, Andy and Rowan Hooper (2002). Darwin and the first ecological experiment. Science, 295, 639-640.
Kohn, David (1980). Theories to work by: rejected theories, reproduction, and Darwin’s path to natural selection. Studies in History of Biology, 4, 67-170.
Kohn, David (1981). On the origin of the principle of diversity. Science, 213, 1105-08.
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Matthew Patrick (1839). Emigration Fields
Mayr, Ernst (1992). Darwin’s principle of divergence. Journal of the History of Biology, 25, 343-359.
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Pearce, Trevor (2010). “A great complication of circumstances” – Darwin and the economy of nature. Journal of the History of Biology, 43, 493-528.
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Wallace, Alfred R. (1856). Attempts at a natural arrangement of birds. Annals and Magazine of Natural History, 18(ser. 2), 193-216.

Sunday, 17 July 2016

Walther May, 1912. "Darwin und Patrick Matthew." Zoologische Annalen 4: 280-311.

Here's a German review of Patrick Matthew's book On Naval Timber and Arboriculture from 1912.

For those not fluent in German, my translation follows. Highlights are my additions. [Square brackets contain my own comments.] They are not part of May's review:
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Darwin and Patrick Matthew.

By 

Prof. Dr. Walther May, Karlsruhe i. B. 

(With a Portrait.) [omitted] 

"It probably belongs to the essence of each discovery," says Rádl in his history of biological theories, "that it struggles for recognition; it forces its wearer to inform you, to fight for them and not infrequently also to suffer. Thousands of ideas struggle for recognition; but most founder in the souls of the absent-minded listening world. The historian must not close his eyes to this fact; he must seek the truth not only to the throne of public recognition, but wherever people think. Through a game of coincidences, an idea is off and on promoted or suppressed; the historian should not be bribed by this, because his goal is to recognize ideas and not to describe the glory of the world. Even if a discovery sank without a trace in the hassles of opinions, it does not therefore cease to form the subject of historiography."
     These words encourage me to renew the memory of a man, whose idea did not go down without a trace, but was disregarded for almost thirty years, before it was awakened by another, larger, to actual life.
     On April 10, 1860 Darwin wrote to Lyell:

"In last Saturday Gardeners’ Chronicle, a Mr Patrick Matthews publishes long extract from his work on “Naval Timber & Arboriculture” published in 1831, in which he briefly but completely anticipates the theory of Nat. Selection.—12 I have ordered the Book, as some few passages are rather obscure but it, is certainly, I think, a complete but not developed anticipation! Erasmus always said that surely this would be shown to be the case someday. Anyhow one may be excused in not having discovered the fact in a work on “Naval Timber”."
     And three days later he wrote to Hooker:
"Questions of priority so often lead to odious quarrels, that I shd. esteem it a great favour if you would read enclosed. If you think it proper that I shd. send it (& of this there can hardly be question) & if you think it full & ample enough, please alter date to day on which you post it & let that be soon.— The case in G. Chronicle seems a little stronger than in Mr. Matthews book, for the passages are therein scattered in 3 places. But it would be mere hair-splitting to notice that.— If you object to my letter please return it; but I do not expect that you will, but I thought that you would not object to run your eye over it."
     On April 21, 1860 "Gardeners Chronicle" brought the following letter Darwin:
"I have been much interested by Mr. Patrick Matthew’s communication in the Number of your Paper, dated April 7th. I freely acknowledge that Mr. Matthew has anticipated by many years the explanation which I have offered of the origin of species, under the name of natural selection. I think that no one will feel surprised that neither I, nor apparently any other naturalist, had heard of Mr. Matthew’s views, considering how briefly they are given, and that they appeared in the appendix to a work on Naval Timber and Arboriculture. I can do no more than offer my apologies to Mr. Matthew for my entire ignorance of his publication. If another edition of my work is called for, I will insert a notice to the foregoing effect."

[Interestingly, May translated the word "naturalist" used by Darwin above as "Naturforscher" into German. Naturforscher, however, means natural scientist or researcher or explorer. Thus May explicated a connotation of the term naturalist in Darwin's use above, that is easily lost in English.]

     According to Francis Darwin, Matthew was not satisfied by this explanation and complained in November 1860, that an article in the "Saturday Analyst and Leader" was hardly fair in calling Darwin the father of the theory of natural selection, because he himself had published all that Darwin tried to prove more than 29 years ago.
     Darwin also recognized Matthew's claims unreservedly in a letter to Quatrefages of 25 April 1861. "I have," he writes there, "lately read M. Naudin’s paper; but it does not seem to me to anticipate me, as he does not shew how Selection could be applied under nature; but an obscure writer on Forest Trees, in 1830, in Scotland, most expressly & clearly anticipated my views—though he put the case so briefly, that no single person ever noticed the scattered passages in his book."
     Later Darwin found that even Matthew had a predecessor still. "Talking about the Origin," he wrote to Hooker in October 1865, "a Yankee has called my attention to a paper attached to Dr Well’s famous Essay on Dew, which was read in 1813 to Royal Soc. but not printed, in which he applies most distinctly the principle of N. Selection to the races of man.—So poor old Patrick Matthew, is not the first, & he cannot or ought not any longer put on his Title pages “Discoverer of the principle of Natural Selection”!" 
     In the historical sketch, that Darwin prefixed to the later editions of his "Origin of Species," he appreciates the Matthew's merits as follows:
     "In 1831 Mr. Patrick Matthew published his work on 'Naval Timber and Arboriculture,' in which he gives precisely the same view on the origin of species as that (presently to be alluded to) propounded by Mr. Wallace and myself in the 'Linnean Journal,' and as that enlarged in the present volume. Unfortunately the view was given by Mr. Matthew very briefly in scattered passages in an appendix to a work on a different subject, so that it remained unnoticed until Mr. Matthew himself drew attention to it in the 'Gardener's Chronicle,' on April 7th, 1860. The differences of Mr. Matthew's view from mine are not much importance: he seems to consider that the world was nearly depopulated at successive periods, and then re-stocked; and he gives as an alternative, that new forms may be generated "without the presence of any mould or germ of former aggregates." I am not sure that I understand some passages; but it seems that he attributes much influence to the direct action of the conditions of life. He clearly saw, however, the full force of the principle of natural selection."
      In the same historical sketch Darwin says about Owen's priority claims:
     "As far as the mere enunciation of the principle of natural selection is concerned, it is quite immaterial whether or not professor Owen preceded me, for both of us, as shown in this historical sketch, were long ago preceded by Dr Wells and Mr. Matthews."
     Besides these judgments by Darwin of Matthew, I only know of two further in the Darwinian literature. Regarding Darwin's remarks on Matthew in the historical sketch, Samuel Butler wrote, in his book "Evolution, Old & New," in 1879:
      "Nothing could well be more misleading. If Mr. Matthew's view of the origin of species is "precisely the same as that" propounded by Mr. Darwin, it is hard to see how Mr. Darwin can call those of Lamarck and Dr. Erasmus Darwin "erroneous"; for Mr. Matthew's is nothing but an excellent and well-digested summary of the conclusions arrived at by these two writers and by Buffon. If, again, Mr. Darwin is correct in saying that Mr. Matthew "clearly saw the full force of the principle of natural selection," he condemns the view he has himself taken of it in his 'Origin of Species,' for Mr. Darwin has assigned a far more important and very different effect to the fact that the fittest commonly survive in the struggle for existence, than Mr. Matthew has done. Mr. Matthew sees a cause underlying all variations; he takes the most teleological or purposive view of organism that has been taken by any writer (not a theologian) except myself, while Mr. Darwin's view, if not the least teleological, is certainly nearly so, and his confession of inability to detect any general cause underlying variations, leaves, as will appear presently, less than common room for ambiguity."
     Contrary to Butler and agreeing with Darwin, Grant Allen called Patrick Matthew the unconscious discoverer of the principle of natural selection in his Darwin Biography (1888), who applied the selection idea, in his book on naval timber, to the whole of nature, sometimes with the same words as Darwin."
     I was led to concern myself with Matthew and his work through an external circumstance. Prof. Dr. P. Unna in Hamburg sent me, on the occasion of the Darwin anniversary in 1909, a letter of the Hamburg pastor, Dr. H. F. Beneke, whose uncle Alexander Matthew (died on 18 January 1911 at the age of 90) was the son of Patrick Matthews. This letter alluded to the priority of Matthew. I then asked the Pastor Beneke for the book and for some biographical notes about his great uncle and received both in amiable manner. Here are the biographical data:
     "Patrick Matthew, born October 20, 1790, died June 8, 1874, married his cousin Christian Nicol (born December 21, 1791, died October 28, 1857). Both their mothers were sisters, born Duncan. From the Duncans he inherited the Gourdie Hill property, actually with the condition to take the name Duncan, what he did not. (The land is now no longer in the family). We also know the names of his parents John Matthew and Agnes Duncan and his grandparents Patrick Matthew in the Rome property on Tay and Helen Millar Broambrae from Fife, but do not have dates for them.
      He must have married about 1819, as the eldest son Robert was born in 1820 and my uncle in 1821. He first managed both Rome and Gourdie Hill, later the latter alone. His picture, as an old man, shows beautiful, noble and pleasant features.
     According to family tradition, the Matthews are from a sister of Robert Bruce, and they are proud to have very square chin, as R. Bruce used to have; but that will of course be treated more jokingly."
      The work on which Patrick Matthew based his priority claims against Darwin, bears the title: "On naval timber and arboriculture:; with critical notes on authors who have recently treated the subject of planting.' It is published by Adam Black, Edinburg; Longman, Rees, Orme, Brown and Green, London, and published in 1831. The content includes XVI and 391 big octavo pages and is organized as follows:

[list of contents omitted] 

     Matthews book is imbued with the patriotic spirit of the Englishman, who desires world domination of his nation. The means to this is for him the seafaring. In the introduction, he noted that the seafaring was of the greatest importance for the improvement of the species, naval superiority almost synonymous with universal dominion, the mainland only the footstool of the mistress of the seas. The periodic recurrence of war seems to him indispensable for the heroic, chivalrous character and the love of freedom. Conflict and fighting should rub the rust from the customs and institutions of his people, the ennobling appeal of danger should arouse the noble passions and the youth be led to emulate the Romans in patriotic thirst for fame, the Spartans in devotion, their own ancestor in bold bravery. Without seeking war, but in preparation to face an enemy, in perhaps not so distant a time, England should maintain its military virtues without malignant sentimentality, especially that which must make up the field of his fame, his navy and their building materials, naval timber.
      Here, it is already indicated that the struggle among nations serves their refinement, but without pronouncing the principle of selection.

[May skipped the second part, probably because it is about ship-building and contains no passage relevant to natural selection.]

     In the third part of the book artificial selection is an issue and natural selection is also hinted at. Since the luxuriance and size of timber is highly dependent on the peculiar variety of species, on the treatment of the seed prior to sowing, and on the treatment of the young plant, and since this fundamental issue is neither much appreciated nor widely understood, Matthew wants to discuss it from scratch. He speaks of the consequences of our lamentable ignorance of the most undeniable facts of natural history: that both the plants and the animals are subjected to an almost unlimited variability in general, due to the climate, the soil, the food and new blending of already formed varieties. In such species, with which man is very familiar, he had become acquainted with these facts, that is, in man himself, the dog, horse, cattle, sheep, poultry, apple, pear, plum, gooseberry, potato and pea, that feature endless varieties, by differing considerably in size, color, taste, firmness of texture, growing season and any recognizable property. In all these species, man seeks to avoid deterioration by careful selection of the biggest and most valuable for further growth, but with timber trees, the inverse procedure was followed. The tall-growing varieties were often cut off prematurely, because of their late seed production, small-growing and weak varieties, in which seed production takes place early and abundantly, on the other hand, had constantly been selected to reproduce because of the ease and convenience with which seeds can be obtained. "May we, then, wonder," asks Matthew, "that our plantations are occupied by a sickly short-lived puny race, incapable of supporting existence in situations where t heir own kind had formerly flourished—particularly evinced in the genus Pinus, more particularly in the species Scots fir; so much inferior to those of Nature's own rearing, where only the stronger, more hardy, soil-suited varieties can struggle forward to maturity and reproduction?"
     This last sentence clearly stated that a selection takes place in nature, leading to the perfection of the race, but the emergence of new species by natural selection is not alleged here. 
     Matthew demands that the farmer gives as much attention to the breeding of his forest trees as to that of his horses, cows and sheep, that he only sows seeds collected from the largest, healthiest and most lavishly growing trees and desists from sowing seeds of precocious or even of the very old and overripe trees, because, by analogy with animals, a weak and early decaying progeny is to be expected from them.
     In the fourth part of his book Matthew says that the benefit of endless seed varieties in the families of plants, even those under nature, probably lies in the fact that one individual (the strongest and best circumstances suited) gains the superiority over others, surpassing and suppressing them, creating space for its full extension and thus at the same time accomplishing a consistent selection of the strongest, best adapted to reproduce. The intervention of man has increased the diversity of varieties, regardless of the new conditions to which he introduced them, by preventing the natural process of selection among the plants, particularly in the more domesticated species, and even in man himself the greater equality and larger strength of the wild tribes can be ascribed to a similar law of selection, in which the weaker individual perishes under the poor treatment on the part of the stronger or from general depression.
     Again, natural selection is considered only as a means of racial improvement and not as a cause of the emergence of new species. 
     Matthew clearly expressed the principle of natural selection as an adapting and perfecting principle in Note B of the Appendix ,"There is," he says here, "a law universal in nature, tending to render every reproductive being the best possibly suited to its condition that its kind, or that organized matter, is susceptible of, which appears intended to model the the physical and mental or instinctive powers, to their highest perfection, and to continue them so. This law sustains the lion in his strength, the hare in his swiftness, and the fox in his wiles. As Nature, in all her modifications of life, has a power of increase far beyond what is needed to supply the place of what falls by Time's decay, those individuals who possess not the requisite strength, swiftness, hardihood, or cunning, fall prematurely without reproducing—either a prey to their natural devourers, or sinking under disease, generally induced by want of nourishment, their place being occupied by the more perfect of their own kind, who are pressing on the means of subsistence."
     Matthew continues this discourse with an argument against hereditary nobility and the law of entail, which appears to him as a mockery of the law of selection, that nature will not let unavenged. He referred to the role of hereditary nobility played in France, the Iberian and Italian peninsula and the Italian islands, and calls on the apologists of the hereditary nobility, primogeniture and the law of entail to say what these countries could have been without the corrupting influence of these unnatural morals. He sees intermittent mixing of the nobility with the people as the only way to protect those against degeneration. In some countries, this mixing would not be necessary as often as as in others, and Britain could be considered as the ground on which the nobility can remain unspoiled the longest. Approaching the equator, however, the renewal would often be necessary, except in high altitudes, in many places every third generation. The repeal of the law of inheritance and primogeniture would increase not only the happiness of the owner in the present state of civilization, increase mortality and give the social order greater strength, but also give the hard work and progress a general incentive whereby the living conditions of the working class would be improved.     
     Even Darwin has called primogeniture a means that was contrary to the action of natural selection. "Our aristocracy," he once wrote to Wallace, "is handsomer (more hideous according to a Chinese or Negro) than middle classes from pick of women; but oh what a scheme is primogeniture for destroying N. Selection.And in the fifth chapter of the "Descent of Man" he discussed the deleterious effects of this system in greater detail, but without neglecting the balancing factors.
     In note C of the Appendix, Matthew examines the causes which have brought about the superiority of a part of the Caucasian race. He finds them mainly in repeated change of place under favorable circumstances. "There are few countries," he says, "where the old breed has not again and again sunk before the vigour of new immigration; we even see the worn out breed, chased from their homes to new location, return, after a time, superior to their former vanquishers, or gradually work their way back in peace, by superior subsisting power: this is visible in France, where the aboriginal sallow Kelt, distinguished by high satyr-like feature, deep-placed sparkling brown or grey eye, narrowed lower part of the face, short erect vertebral column, great mental acuteness, and restless vivacity, has emerged from the holes of the earth, the recesses of the forests and wastes, into which it had been swept before the more powerful blue-eyed Caucasian; and being a smaller, more easily subsisting animal, has, by starving and eating out, been gradually undermining the breed of its former conquerors."
     But even more than the change of location the related circumstances have their share in the perfection of the species. In the unrest that accompanied the emigration, the varieties strongest in mind and body take over the leader role, impressing their character onto the people at large and constituting the reproductive part, while the weaker varieties generally perish at the occasional hardships. When a cohort emigrates from a community, it will generally consist of the bolder and brisked spirits, who will use the right of the conquerors to connect with the best of the natives that they overwhelm; their choice among these will be determined by personal characteristics and not by the accidental circumstances of wealth or high birth—a consideration which leads to the degradation of the race and one of the reasons, why the nobility of Europe is so inferior in comparison with the Asian. 
     Again, selection is only recognized as a means of racial improvement.
     In note D Matthew repeats the thought, already indicated in the introduction, that national power and size is impossible without the operation of the egotistical drives. Our milder manners, our benevolence our tranquility, kindness and sense of refinement our sweet dreams of peace and joy, he calls a negative weight in the scales of national strength. The stronger excitation of hatred, ambition, pride, patriotism and more selfish passions, he deems necessary for the full and strong development of national energy. That Britain had impressed its ability and morality to a considerable extent onto the world, is due to the fact that it first ravaged these countries with fire and sword. 
     These words remind us of the consequences that have drawn some modern racial theorists and many anti-darwinian ethicists from Darwinism.

[May skipped the end-notes E and F of the appendix. Note E was on the improper calculation of register tonnage of trading vessels leading to ill constructions of the vessels, in order to safe charges for lights, harbours and other dues. Note F was on the geological history of the North Sea, which Matthew called the German Ocean. Next May refers to a "long epilogue." This is actually a colophon following after the end of Note F and a horizontal line indicating this. Whereas footnotes in the main text refer to the end-notes A-F of the appendix, the main text contains no reference to this colophon suggesting that it was an afterthought.

     Matthew concludes his work with a long epilogue, in which he set out his understanding of the evolution of organic life on Earth. Only here, he regarded natural selection not only as a means of racial improvement and adaptation, but also as a cause of the origin of new species and the organic development at all. He speaks, first, of a power of change under a change of circumstances, which belongs to the living substance, or rather the hodgepodge of low life, which seems to form the higher, and that one must admit, if one does not want to accept a repeated wonderful creation. The changes that have happened as a result of human intervention with the domestic animals and crops before him, and the likelihood that the living conditions were very different in the various geological periods, but consistent within each, seem to prove to him the accuracy of the assumption that, at the beginning of each new age [meaning geological eras between catastrophes], no creation took place, but some organisms surviving from the former age have, again, adapted their existence over time to the change of circumstances and to every possible kind of living conditions. "Is the inference, then, unphilosophic," he asks, in consideration of the large chemical changes of the water and the atmosphere, "that living things, which are proved to have a circumstance-suiting power—a very slight change of circumstances by inducing a corresponding change of character—may have gradually accommodated themselves to the variations of the elements containing them, and, without new creation, have presented the diverging changeable phenomena of past and present organized existence."
     Matthew evidently connects in his somewhat vague hypothesis the doctrine of catastrophism with the theory of evolution. He believes that the destructive liquid streams, which separated the ages and destroyed almost all living things, reducing being so much, that an unoccupied field got available for new diverging branches of life adapting itself to the new conditions and then, after completion of this adjustment, remained constant for the duration of their existence, except for the few residues that reached alive into the next following period. 
     Besides this theory of evolution and the doctrine of creation there is, according to Matthew, only one alternative explanation of organic changes, namely the assumption of an "indestructible or molecular life, gradually uniting and developing itself into new circumstance-suited living aggregates, without the presence of any mould or germ of former aggregates," which probably means the re-emergence of life by spontaneous generation, which Matthew distinguishes from new creation only by the fact that it "forms a portion of a continued scheme or system."
     Thus, Matthew has taken his position concerning the three possible hypotheses of biogenesis, on creation, on spontaneous generation and on evolution, and he opted for the theory of evolution; but it can hardly be called an advance over the earlier development theorists such as Buffon, Erasmus Darwin and Lamarck, when he connects it to the doctrine of catastrophism.
     Far more important than the views of Matthews on organic change in general are his ideas about the causes of these changes. Here he raises the following question: "Do they arise from admixture of species nearly allied producing intermediate species? Are they the diverging ramifications of the living principle under modification of circumstance? Or have they resulted from the combined agency of both? Is there only one living principle? Does organized existence, and perhaps all material existence, consist of one Proteus principle of life capable of gradual circumstance-suited modifications and aggregations?"
     Matthew cannot accept that much of the changes in the organisms is owed to the mixing of closely related species, since all change is very limited by this, and limited in the circle of what is called species. He sees the main cause of change in the self-regulating adaptability of organisms, which he ascribes, at least partially, to the immense fertility of nature.
    "Nature, who, as before stated, has, in all the varieties of her offspring," he says, "a prolific power much beyond (in many cases a thousandfold) what is necessary to fill up the vacancies caused by senile decay. As the field of existence is limited and pre-occupied, it is only the hardier, more robust, better suited to circumstance individuals, who are able to struggle forward to maturity, these inhabiting only the situations to which they have superior adaptation and greater power of occupancy than any other kind; the weaker, less circumstance-suited, being prematurely destroyed. This principle is in constant action, it regulates the colour, the figure, the capacities, and instincts; those individuals of each species, whose colour and covering are best suited to concealment or protection
from enemies, or defence from vicissitude and inclemencies of climate, whose figure is best accommodated to health, strength, defence, and support; whose capacities and instincts can best regulate the physical energies to self-advantage according to circumstances—in such immense waste of primary and youthful life, those only come forward to maturity from the strict ordeal by which
Nature tests their adaptation to her standard of perfection and fitness to continue their kind by reproduction.
      From the unremitting operation of this law acting in concert with the tendency which the progeny have to take the more particular qualities of the parents, together with the connected sexual system in vegetables, and instinctive limitation to its own kind in animals, a considerable uniformity of figure, colour, and character, is induced, constituting species; the breed gradually acquiring the very best possible adaptation of these to its condition which it is susceptible of, and when alteration of circumstance occurs, thus changing in character to suit these as far as its nature is susceptible of change."
     In these sentences, the principle of natural selection is expressed with full clarity and widely applied, and Darwin is certainly right when he says of Matthew "He clearly saw the full implications of the principle of natural selection." It is impossible to see how Butler can discard this interpretation of Matthew's train of thought, and claim that the Scottish writer has only represented the doctrines of Buffon and Erasmus Darwin. The principle of natural selection is there, although Matthew also added that this adaptive law does not exclude the influence of the will or feeling on the design of the body. Matthew just recognizes other factors of species transformation in addition to natural selection, exactly like Darwin did, on whose scope further research had to decide. "to investigate," he says, "how much variation is modified by the mind or nervous sensation of the parents, or of the living thing itself during its progress to maturity; how far it depends upon external circumstance, and how far on the will, irritability and muscular exertion, is open to examination and experiment. In the first place, we ought to investigate its dependency upon the preceding links of the particular chain of life, variety being often merely types or approximations of former parentage; thence the variation of the family, as well as of the individual, must be embraced by our experiments."
     That is, Matthew already advocated the most modern direction of the development theory, the experimental.
      In the further course of his argument, he stressed that the continuation of the family type is both physical and mental, and is evidenced by many of the dispositions or instincts of the different human races. He regarded these native or inherited ideas or habits, that prevail especially in insects, as an "abiding memory" and believed to solve a lot of the mystery of instinct and the foreknowledge that these animals have of what is necessary, in order to complete their round of life, by reducing the instincts to knowledge, or impressions, and habits, acquired by a long experience. So he explains, here, the instincts according to Lamarck's principle; that he also subjects them to natural selection, however, is clear from his earlier statements.
     In discussing the instincts of insects, Matthew also touches on the problem of individuality. He finds it difficult to determine the specific points, in some insects, when each individuality begins with the different stages of egg, larva, pupa, or whether much consciousness of individuality exists.
     The epilogue concludes with a reflection on the imbalance of nature introduced by humans. They have in the present age attained a mastery of the material world and a successful power of multiplication, which makes it likely that the whole surface of the earth will soon be overrun by this engrossing anomaly, to the annihilation of every wonderful and beautiful variety of animated existence, that does not administer the human needs.
     It is admirable how many problems are touched in Matthew's book. However, only the attempt of a comprehensive application of the principle of selection on the whole organic nature is fundamentally new. All the other ideas of Matthew had already been expressed by earlier thinkers. Even the principle of natural selection as such had already been pronounced by Wells and Prichard, but they applied it to the races of man only. In comparison with this very limited application, Matthew's hypothesis seems to be a new thought. We can therefore call the Scottish landowner the first, who recognized natural selection as a general principle of nature, without scruple. But he does not seem to have considered this principle to be as important for biological science, as Darwin later adjudged it, otherwise he would not have published his ideas in the appendix to a work on naval timber. But even if he had published this idea, in the existing form, in a separate document, it would have had as little influence on the science, as had the short treatises that Wallace and Darwin published in 1858 on the theory of natural selection in the Journal of the Linnaean Society. Because, as Butler rightly remarks, the same reproach must be made against Matthew's abstract of the theory of evolution as to Erasmus Darwin's view of this theory, that it was in fact too short. "It may be true," says Butler, "that brevity is the the soul of the wit, but the leaders of science will generally succeed in burking new-born wit, unless the brevity of its soul is found compatible with a body of some bulk."
     Darwin first gave the body to the soul of the theory of natural selection. The ingenious way in which he related the facts of almost all biological disciplines in terms of the selection idea into an organic whole, is his very own merit, that defies all other priority claims.   

Literature.
Allen, Qraot, Charles Darwin. London, Longmans, Green, and Co. 1888.

Butter, Samuel, Evolution, old and new. London, Hardwicke and Bogue, 1879.

Darwin, Charles, Die Entstehung der Arten. Deutsch von Victor Carus. 8. Auflage. Stuttgart, E. Schweizerbart, 1899.

Darwin, Francis, Leben und Briefe von Charles Darwin. Deutsch von Victor Carus. 2. Auflage. Stuttgart, E. Schweizerbart, 1899.

Darwin, Francis and A. C. Seward, More letters of Charles Darwin. London, John Murray, 1903.

Matthew, Patrick, On naval timber and arboriculture; with critical notes on authors who have recently treated the subject of planting. Edinburgh, Adam Black; London, Longman, Rees, Orme, Brown and Green, 1831.

Rádl, Em., Geschichte der biologischen Theorien. II. Teil. Leipzig, Wilhelm Engelmann, 1909.