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Origin of Species
chapter iii. struggle for existence   Its bearing on natural selection
Charles Darwin
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       Before entering on the subject of this chapter I must make a few preliminary remarks to show how the struggle for existence bears on natural selection. It has been seen in the last chapter that among organic beings in a state of nature there is some individual variability: indeed I am not aware that this has ever been disputed. It is immaterial for us whether a multitude of doubtful forms be called species or sub-species or varieties; what rank, for instance, the two or three hundred doubtful forms of British plants are entitled to hold, if the existence of any well-marked varieties be admitted. But the mere existence of individual variability and of some few well-marked varieties, though necessary as the foundation for the work, helps us but little in understanding how species arise in nature. How have all those exquisite adaptations of one part of the organisation to another part, and to the conditions of life and of one organic being to another being, been perfected? We see these beautiful co-adaptations most plainly in the woodpecker and the mistletoe; and only a little less plainly in the humblest parasite which clings to the hairs of a quadruped or feathers of a bird; in the structure of the beetle which dives through the water; in the plumed seed which is wafted by the gentlest breeze; in short, we see beautiful adaptations everywhere and in every part of the organic world.
       Again, it may be asked, how is it that varieties, which I have called incipient species, become ultimately converted into good and distinct species, which in most cases obviously differ from each other far more than do the varieties of the same species? How do those groups of species, which constitute what are called distinct genera and which differ from each other more than do the species of the same genus, arise? All these results, as we shall more fully see in the next chapter, follow from the struggle for life. Owing to this struggle, variations, however slight and from whatever cause proceeding, if they be in any degree profitable to the individuals of a species, in their infinitely complex relations to other organic beings and to their physical conditions of life, will tend to the preservation of such individuals, and will generally be inherited by the offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. I have called this principle, by which each slight variation, if useful, is preserved, by the term natural selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art.
       We will now discuss in a little more detail the struggle for existence. In my future work this subject will be treated, as it well deserves, at greater length. The elder De Candolle and Lyell have largely and philosophically shown that all organic beings are exposed to severe competition. In regard to plants, no one has treated this subject with more spirit and ability than W. Herbert, Dean of Manchester, evidently the result of his great horticultural knowledge. Nothing is easier than to admit in words the truth of the universal struggle for life, or more difficult--at least I found it so--than constantly to bear this conclusion in mind. Yet unless it be thoroughly engrained in the mind, the whole economy of nature, with every fact on distribution, rarity, abundance, extinction, and variation, will be dimly seen or quite misunderstood. We behold the face of nature bright with gladness, we often see superabundance of food; we do not see or we forget that the birds which are idly singing round us mostly live on insects or seeds, and are thus constantly destroying life; or we forget how largely these songsters, or their eggs, or their nestlings, are destroyed by birds and beasts of prey; we do not always bear in mind, that, though food may be now superabundant, it is not so at all seasons of each recurring year.
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Introduction
chapter i. variation under domestication
   Causes of Variability
   Effects of Habit and the use or disuse of Parts; Correlated Variation; Inheritance
   Character of Domestic Varieties; Difficulty of distinguishing between Varieties and Species; Origin of Domestic Varieties from one or more Species
   Breeds of the Domestic Pigeon, Their Differences and Origin
   Principles of Selection, anciently followed, their Effects
   Unconscious Selection
   Circumstances favourable to Man's power of Selection
chapter ii. variation under nature
   Variability
   Individual Differences
   Doubtful species
   Wide ranging, much diffused, and common species, vary most
   Species of the larger genera in each country vary more frequently than the species of the smaller genera
   Many of the species of the larger genera resemble varieties in being very closely, but unequally, related to each other, and in having restricted ranges.
   Summary
chapter iii. struggle for existence
   Its bearing on natural selection
   The term, Struggle for Existence, used in a large sense
   Geometrical ratio of increase
   Nature of the checks to increase
   Complex relations of all animals and plants to each other in the struggle for existence
   Struggle for life most severe between individuals and varieties of the same species
chapter iv. natural selection; or the survival of the fittest
   Natural Selection
   Sexual Selection
   Illustrations of the action of Natural Selection, or the survival of the fittest
   On the Intercrossing of Individuals
   Circumstances favourable for the production of new forms through Natural Selection
   Extinction caused by Natural Selection
   Divergence of Character
   The Probable Effects of the Action of Natural Selection through Divergence of Character and Extinction, on the Descendants of a Common Ancestor
   On the degree to which Organisation tends to advance
   Convergence of character
   Summary
chapter v. laws of variation
   Effects of changed conditions
   Effects of the increased use and disuse of parts, as controlled by Natural Selection
   Acclimatisation
   Correlated variation
   Compensation and economy of growth
   Multiple, rudimentary, and lowly organised structures are variable
   A part developed in any species in an extraordinary degree or manner, in comparison with the same part in allied species, tends to be highly variable
   Specific characters more variable than generic characters
   Secondary sexual characters variable
   Distinct species present analogous variations, so that a variety of one species often assumes a character proper to an allied species, or reverts to some of the characters of an early progenitor
   Summary
chapter vi. difficulties of the theory
   Difficulties of the theory of descent with modification
   Absence or rarity of transitional varieties
   On the origin and transition of organic beings with peculiar habits and structure
   Organs of extreme perfection and complication
   Modes of transition
   Special difficulties of the theory of Natural Selection
   Organs of little apparent importance, as affected by Natural Selection
   Utilitarian doctrine, how far true: Beauty, how acquired
   Summary
chapter vii
   Miscellaneous Objections to the Theory of Natural Selection
chapter viii. instinct
   Instincts comparable with habits, but different in their origin
   Inherited changes of habit or instinct in domesticated animals
   Special instincts; Instincts of the cuckoo
   Slave-making instinct
   Cell-making instinct of the hive-bee
   Objections to the theory of natural selection as applied to instincts: neuter and sterile insects
   Summary
chapter ix. hybridism
   Distinction between the sterility of first crosses and of hybrids
   Degrees of sterility
   Laws governing the sterility of first crosses and of hybrids
   Origin and causes of the sterility of first crosses and of hybrids
   Reciprocal dimorphism and trimorphism
   Fertility of varieties when crossed and of their mongrel offspring not universal
   Hybrids and mongrels compared independently of their fertility
   Summary of Chapter
chapter x. on the imperfection of the geological record
   On the absence of intermediate varieties at the present day
   On the lapse of time, as inferred from the rate of denudation and of deposition
   On the poorness of our palaeontological collections
   On the absence of numerous intermediate varieties in any single formation
   On the sudden appearance of whole groups of allied species
   On the sudden appearance of groups of allied species in the lowest known fossiliferous strata
chapter xi. on the geological succession of organic beings
   On the slow and successive appearance of new species
   On extinction
   On the forms of life changing almost simultaneously throughout the world
   On the affinities of extinct species to each other and to living species
   On the state of development of ancient compared with living forms
   On the succession of the same types within the same areas, during the later Tertiary Periods.
   Summary of preceding and present chapter
chapter xii. geographical distribution
   Present distribution cannot be accounted for by differences in physical conditions
   Single centres of supposed creation
   Means of dispersal
   Dispersal during the Glacial period
   Alternate Glacial periods in the north and south
chapter xiii. geographical distribution -- continued
   Distribution of fresh-water productions
   On the inhabitants of oceanic islands
   Absence of Batrachians and terrestrial Mammals on oceanic islands
   On the relation of the inhabitants of islands to those of the nearest mainland
   Summary of the last and present chapter
chapter xiv. mutual affinities of organic beings: morphology -- embryology -- rudimentary organs
   Classification
   Analogical resemblances
   On the nature of the affinities connecting organic beings
   Morphology
   Development and embryology
   Rudimentary, atrophied, and aborted organs
   Summary
chapter xv
   Recapitulation and Conclusion
Glossary of Scientific Terms