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I cannot grasp the structure of this sentence.

Natural selection will tend to maximize r for the environment in which the species lives, for any mutation or gene combination which increases the chance of genotypes possessing them contributing more individuals to the next generation (that is, of increasing r) will be selected over genotypes contributing fewer of their kind to successive generations. (Birch 1960, p. 10).

(r is rate of increase.)

  1. Is "for" after comma is in parallel with former "for" after "maximize r" or just a conjunction like "because" ?

  2. What is the meaning of "possessing them" here?

  3. Why could "possessing" and "contributing" be appeared in succession ?

  4. What (is the subject of) "will be selected over"?

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    The string in bold looks like a reason adjunct where "for" has the same meaning as "because". I'm not sure if "them" is anaphoric to "the species" or some previously mentioned noun, so "possessing them" means "possessing x". "Genotypes possessing them" is subject of "contributing more individuals ...". The subject of "will be selected over" is "any mutation or gene combination which increases the chance of genotypes possessing them contributing more individuals to the next generation (that is, of increasing r)". I think!! – BillJ Mar 26 '17 at 8:43
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    @Bill - Academic writing is often hard to understand on first reading and may well be intended that way. Building on Bill's good job, I read "possessing them" to refer to "any mutation or gene combination." The genotypes possessing mutations contributing more to the next generation "will be selected." In the original version, "any mutation or gene combination" is the subject of "will be selected." – Yosef Baskin Mar 28 '17 at 19:28
  • The second "for" is comparable to because. The sentence becomes difficult because "contributing more individuals to the next generation (that is, of increasing r)" is parenthetical and should be enclosed in commas. – Zan700 Dec 12 '18 at 13:52
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  1. No, "for" means 'for the following reason, because' in this sentence.
  2. The expression "possessing them" is a reduced relative clause modifying "genotypes". The pronoun "them" refers to "mutation or gene combination". It means 'those genotypes (~set of all genes) that posesses a particular mutation or gene combination (~a particular gene)'.
  3. The verb "possessing" modifies "genotypes", as described under 2. The verb "contributing" complements "chances of". The quote would be easier to understand if it read, "increase the chance of genotypes possessing them to contribute more individuals". The subject of "contributing" is "genotypes possessing them." The functions of "possessing" and "contributing" are completely different. Hence there is nothing that would prevent the two items from co-occurring in succession.
  4. The subject of "will be selected over" is "any mutation or gene combination which increases the chance of genotypes possessing them contributing more individuals to the next generation (that is, of increasing r)", or, simplified, "any mutation or gene combination [...] will be selected over".

The quote can be paraphrased as follows:
The reason is as follows (for):
There are mutations or gene combinations.
Certain genotypes may possess such mutations or gene combinations (genotypes possessing them).
Such genotypes may have a higher chance of contributing individuals to the next generation (chances of ... contributing individuals).
Hence, these mutations or gene combinations are selected.

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