Wednesday, August 14, 2013

Lamarckism Revisited

  The concept that traits acquired during an individual's lifetime can be passed on to offspring and therefore affect evolution—a theory most associated with the French scientist Jean-Baptiste Lamarck (1744-1829)—was relegated to the dust bin by the advent of Darwinism.  After Darwin, a new orthodoxy prevailed for more than a century.  In present-day terminology, it held that only traits already embedded in genes are passed on, and evolution occurs exclusively by natural selection of mutations in the genome.  Such mutations are purely random events, caused by errors in DNA transcription or by environmental factors.  Any suggestion of "directed" mutations—beneficial, nonrandom changes in the genome provoked by an organism in response to specific environmental challenges—was decried.  Further, the notion that changes in and inheritance of traits could occur through other than genetic means was anathema.

  That orthodox Darwinian viewpoint has been challenged in the past 30 years or so.  Modern studies have shown that directed, nonrandom genomic mutations can occur; that acquired non-genomic attributes can be inherited; and that both are subject to natural selection in the same way as are random genomic mutations.  These new insights and much else are discussed in a remarkable, but lengthy and hard-to-read book by Eva Jablonka and Marion J. Lamb, Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral and Symbolic Variation in the History of Life.

  I will skip over Jablonka-Lamb's first dimension, genetics, for it is the Darwinian dimension, with which I assume you are familiar.  The second dimension, epigenetics, is the most astounding to me.  It turns out that molecular "markers" can attach themselves to the genome as a result of environmental impact or stress.  They do not change the genome, but can turn genes on and off, thus changing the expression of an individual's genetic traits.  What is surprising is that these epigenetic factors can be passed on to offspring.

  The earliest-discovered and most common epigenetic marker is methylation of DNA.  Small methyl (CH3) groups can become attached to bases in the DNA sequence because of environmental influences and stress, but they do not change the DNA other than by turning genes on or off.  Remarkably, the methylation remains in daughter cells after cell division.  Even more remarkably, when methylation affects sperm or egg cells, it is inherited by an embryo and thus passed on to the next generation.  To quote from Jablonka-Lamb:

"[B]ecause it provides an additional source of variation, evolution can occur through the epigenetic dimension of heredity even if nothing is happening in the genetic dimension.  But it means more than this.  Epigenetic variations are generated at a higher rate than genetic ones, especially in changed environmental conditions, and several epigenetic variations may occur at the same time.  Furthermore, they may not be blind to function, because changes in epigenetic marks probably occur preferentially on genes that are induced to be active by new conditions.  This does not mean that all induced changes are adaptive, but it does increase the chances that a variation will be beneficial.  The combination of these two properties—a high rate of generation and a good chance of being appropriate—means that adaptation through the selection of epigenetic variants may be quite rapid compared with adaptation through genetic change."

This one astonishing paragraph was to me worth the price of the book.

  Less compelling to me, but still impressive, are Jablonka and Lamb's asserted third and fourth dimensions of evolution—behavioral and symbolic.  If I understand these dimensions correctly, they overlap with and extend what the British evolutionary biologist Richard Dawkins called "memes"—an analog of genes.  Just as a gene carries information about biological traits, a meme carries information about socio-cultural ideas, symbols, and practices that are passed from generation to generation but not necessarily to direct offspring.  That many memes are acquired under the influence of the environment and unrelated individuals, and propagated by social interaction rather than genetics, doesn't sound extraordinary—we know that species, especially Homo sapiens, pass on a vast repertoire of behaviors derived from their environment and social groupings.  More startling, however, is evidence Jablonka and Lamb cite indicating that acquired memes may also be inherited through bio-chemical means; for example, a mother's particular food preferences have been shown to be reproduced in her offspring through placental affects on the fetus.  The important point Jablonka and Lamb make is that memes as well as genes are subject to evolutionary selection: those that are advantageous for survival will persist, those that aren't will vanish.

  The focus of the Jablonka-Lamb book is therefore on information transmission as the basis of evolution, not only the biological information of the genome and its epigenetic markers, but also the behavioral and symbolic information of cultures. 

"All four ways of transmitting information introduce, to different degrees and in different ways, instructive mechanisms into evolution.  All shape evolutionary change. …  As molecular biology uncovers more and more about epigenetic and genetic inheritance, and as behavioral studies show how much information is passed on to others by nongenetic means, evolutionary biologists will have to abandon their present concept of heredity, which was fashioned in the early days of genetics."

For those of us who have been raised in the pure-genetics tradition, this is a big gob to swallow.  We were taught to ignore Lamarck and to disdain the USSR's ideologically motivated Lysenko;  now, it seems, some of their ideas may turn out to have merit.