Haeckel had a point
My colleague Paul Strode wrote a very clear and concise explanation of Ernst Haeckel's "ontogeny recapitulates phylogeny" law for our book Why Evolution Works (and Creationism Fails). In Chapter 11, Strode explains that Haeckel was wrong in thinking that embryos resemble the ancestral adult forms; rather, early embryos resemble the embryos of ancestral forms. In other words, Haeckel was on to something, but he didn't get it quite right. Strode explains further, "Recapitulation nevertheless provides helpful insight into evolutionary relationships and ancestry," and argues that von Baer's law is closer to the truth. Chapter 11 follows:
In 1874, a German comparative embryologist and Darwin enthusiast, Ernst Haeckel, began publishing his drawings of vertebrate embryos in various stages of development to explain common ancestry and support Darwin's theory of evolution. Haeckel drew the first phylogenies and coined the term "tree of life." His skillful and detailed drawings clearly showed that the early-stage embryos of vertebrates are nearly if not wholly identical. With his drawings and accompanying descriptions, Haeckel promoted the idea that "ontogeny recapitulates phylogeny," also called the biogenic law. The biogenic law states that the path of an organism during its embryological development (ontogeny) is a summary of its evolutionary history (phylogeny). For example, a human embryo has structures that resemble gill slits, just like a fish. The human embryo then loses the gill slits and grows a tail and four limbs, resembling a reptile. The tail disappears, and the embryo begins resembling a primate. Haeckel tirelessly promoted his biogenic law, and many biologists in the nineteenth and into the twentieth centuries embraced it as a simple explanation of evolution. How convenient that all we had to do was look at embryological development to understand an organism's evolutionary history!
While Haeckel had many supporters, he was not without his early critics. Almost immediately, many of Haeckel's fellow embryologists noticed that he had taken artistic liberties in his drawings to support his ideas, yet few rejected the biogenic law outright. One of the first written criticisms of the biogenic law appeared in an 1894 article by zoologist Adam Sedgwick in the Quarterly Journal of Microscopic Science. Sedgwick argued that the biogenic law conflicted with a principle known as von Baer's law, after Karl Ernst von Baer, one of the founders of embryology. Von Baer had noted, contrary to Haeckel, that the embryos of higher animal forms resembled the embryos, not the adults, of earlier forms. Thus, for example, a human embryo may pass through a stage where it resembles a fish embryo, but not an adult fish. Sedgwick noted, "Embryos of different members of the same group are more alike than the adults, and the resemblances are greater the younger the embryos examined." He continued that when the actual embryos were examined "a blind man could distinguish between them."
As the twentieth century unfolded and empirical embryology and genetics emerged, it became clear that Haeckel had emphasized similarities between the embryos of various vertebrate classes in his drawings and neglected the differences. It is not clear whether he purposely altered his drawings to better fit his ideas. Even so, the drawings fascinated lay people and scientists outside the fields of embryology and evolution. Biology textbook authors looking for images to illustrate their chapter pages happily included the drawings. Due to the cost-effective practice of recycling images and accompanying explanations, as well as the paucity or even lack of evolutionary biologists on the editorial staffs of textbook companies, the drawings remained in many textbooks until the 1970s. During this time, many teachers who had little or no background in evolution taught their students that the drawings were evidence of evolution, and the students were encouraged to understand and memorize the catchy phrase, "ontogeny recapitulates phylogeny."
Finally, in 1977, in his technical book Ontogeny and Phylogeny, Stephen Jay Gould carefully dissected Haeckel's drawings and disproved the general ideas behind the biogenic law. Later, in one of his last essays in Natural History magazine (March 2000), Gould explained that while Ernst Haeckel was regarded among his contemporaries as a master naturalist, he often "took systematic license in 'improving' his specimens to make them more symmetrical or more beautiful." Most likely as a result of Gould's careful critique of Haeckel, Haeckel's drawings are no longer found in today's biology textbooks.
Haeckel was not wholly wrong. As we will see below, phylogeny is to some extent recapitulated in embryological development, but it does not recapitulate the adult forms. Rather, embryological development of today's vertebrates summarizes the evolution of past embryos, not past adults. Haeckel's conception that evolution is inherently progressive, however, is no longer accepted. Whether Haeckel deliberately fudged his drawings remains a matter of debate. Unfortunately, the creationist Jonathan Wells, in the book Icons of Evolution [discussed in Chapter 3], exaggerates the importance of Haeckel and the biogenic law, and purports to show that the entire theory of evolution is founded on a handful of errors such as the biogenic law. In reality, as we have seen, biologists disputed the biogenic law early on and eventually replaced it with the more-realistic von Baer's law. The eventual discovery of Haeckel's misstep is an excellent example of how science self-corrects. That such mistakes are uncovered is not a weakness of science, but a strength. Haeckel's drawings were mere bumps on the road to a comprehensive theory of evolution.
Recapitulation
In its original form, the idea that "ontogeny recapitulates phylogeny" is not an accurate evolutionary statement. Recapitulation nevertheless provides helpful insight into evolutionary relationships and ancestry. Harvard University zoologist Ernst Mayr described recapitulation in embryological development as the appearance of an ancestral structure that is found in two different lineages, say, for example, the pharyngeal arches and pouches (see below) that are found in the embryos of both fish and mammals. The structure then disappears from the embryo (or provides the organizational foundation for future structures) in one lineage, mammals, but is maintained in the adult form of the other lineage, fish. Mayr argued that this structure provides evidence that these two lineages are, in fact, connected to a common ancestor, that is, a group of organisms, not a single individual, from which two lineages most likely descended.
But does not an organism waste energy on a structure that will disappear later during embryological development? To the contrary, these ephemeral ancestral features provide a framework upon which the successful development of future structures depends. In other words, these ancestral characteristics are now playing new roles in embryological development, like organizing tissues that will eventually become bones in the skeleton of an individual. For example, the notochord is a character that provides internal structural support and unites all members of the phylum Chordata (or chordates, animals with notochords and pharyngeal arches, among other characteristics). At some point in development, every species within Chordata, from sea squirts to sea lions, possesses a notochord. Most members of the chordate phylum, those in the subphylum Vertebrata, replace the notochord with vertebrae (the backbone) later in development to support the pelvic and pectoral girdles (to which the front and hind limbs are attached) and to protect the spinal cord. Yet in vertebrates, as the notochord is forming (during a developmental process called gastrulation), it first functions to establish the midline of the embryo. Once the notochord forms, its cells send molecular signals to neighboring cells, inducing those cells to begin developing divisions of the central nervous system and eventually the brain and spinal cord. Thus, without the notochord functioning as an organizer in an early vertebrate embryo, the development of the entire central nervous system of the individual is compromised. In the two other subphyla of the chordates, Cephalochordata and Urochordata, the notochord is either maintained in the adult (Cephalochordata such as Lancelets) or present in the larval form and lost during metamorphosis into the adult form (Urochordata such as Tunicates).
Another character that unites the chordates is the pharyngeal arches and pouches (sometimes inaccurately called gill arches and gill slits). It is not advantageous for terrestrial animals with lungs to retain the gills that develop from the pharyngeal arches and pouches in fish species. The successful embryological development of terrestrial vertebrates, however, requires the organizing presence of these early structures. For example, among other things, the arches give rise to facial bones, parts of the inner and outer ears, and the cartilage of the larynx. The pouches give rise to the Eustachian tubes, and the thyroid and thymus glands. Proper interaction between the arches and pouches is required for normal embryological development. For example, a mutation on chromosome 22 in humans allows an improper association between these early structures and results in the thymus failing to develop. The resulting syndrome, DiGeorge syndrome, causes recurrent infections, heart defects, and facial bone malformation.
Conclusion
Whereas Haeckel's recapitulation theory was wrong in detail, recapitulation, as we now understand it, provides powerful support for Darwin's ideas of common ancestry and descent with modification. Because development is genetically controlled, small changes in developmental genes can have substantial implications for an individual's juvenile and adult form and function, and on its eventual ability to survive and produce offspring of its own.
In the next chapter, we will introduce and describe a new field of biology that shows, through the biology of genes, how an individual passes from a single-celled egg to a multi-celled adult and how a fish journeys through evolutionary time to eventually arrive as a human. This is the field of evolutionary developmental biology, or Evo Devo.
I am very sorry, but if you want to read the next chapter, you will have to buy the book.
A similar argument was presented briefly here in the text and the comments.
Chapter 11 of Why Evolution Works (and Creationism Fails), by Matt Young and Paul K. Strode. Copyright © 2009 by Matt Young and Paul K. Strode. All rights reserved. Reprinted with permission.
35 Comments
John Kwok · 20 June 2010
Excellent post, Matt, and thanks for the reminder regarding Strode's excellent summation in that superb volume of yours (which I might add, ought to be on everyone's reading list merely to understand and to defuse cretinous arguments from creos, especially those of the Dishonesty Institute.
James F · 20 June 2010
What amazes me is that this all translates, in creationist terms, to "Haeckel was a fraud, therefore evolution is false."
Ichthyic · 20 June 2010
What amazes me is that this all translates, in creationist terms, to “Haeckel was a fraud, therefore evolution is false.”
why?
the gullible have been falsely claiming fraud at large since before piltdown man, and will continue to do so every time there is the slightest bit of misinformation posted by (and inevitably discovered and corrected by) scientists.
they.
don't.
like.
science.
MrG · 20 June 2010
Ichthyic · 20 June 2010
http://scienceblogs.com/pharyngula/2007/06/return_of_the_son_of_the_bride.php
...just in case anyone was wondering if the Dishonesty Institute has done this before.
MrG · 20 June 2010
Joel · 20 June 2010
Excellent post! Thanks for this!
James F · 20 June 2010
Stanton · 20 June 2010
When such people are asked "why do you hate science?" to which they always reply, "I don't hate science, I just hate how those evil materialistic scientists (ab)use science to brainwash children into becoming evil, God-hating atheists. Oh, and I don't like learning about science because it's of the Devil."
MrG · 20 June 2010
Or you get the "bad science" "good science" game, as in: "I'm not against science, it's just that evolution is bad science."
"So how do you define 'bad science'?"
"That's any science that tells me anything that I don't want to hear."
"And 'good science'?"
"That's all the rest of science that I'm ignoring for the moment until it tells me something I don't want to hear."
John Vanko · 20 June 2010
robert van bakel · 20 June 2010
Italy 1; New Zealand 1; I think I'm going to have to forego my atheism, there must indeed be a god.
Ichthyic · 20 June 2010
...Friday is Paraguay.
Dale Husband · 20 June 2010
fnxtr · 20 June 2010
Dale stop pussyfooting and tell us what you really think... ;-)
Troy Britain · 21 June 2010
Ntrsvic · 21 June 2010
Frank J · 21 June 2010
MrG · 21 June 2010
raven · 21 June 2010
John Kwok · 21 June 2010
Matt Young · 21 June 2010
Matt Young · 21 June 2010
I finally found Hellman's article here, on the website of the parent Jerusalem Post.
raven · 21 June 2010
Dale Husband · 21 June 2010
Matt Young · 21 June 2010
I will allow Mr. Husband's self-described rant to stand, but I consider it very far off task and will limit responses. I will not go into more detail, but I also consider it intemperate and not entirely accurate as regards the founding of Israel.
I do not want to get into an overtly political discussion, but I frankly could not agree more with Mr. raven's comment.
harold · 21 June 2010
Matt Young -
For balance, I will express my views; please feel free to delete them if you wish.
I think that the Israeli education situation described in that article is shocking, and highly instructive for the US.
In some ways the situation here is almost worse, as a major political faction has adopted science denial on a number of issues as part of its defining platform (global warming denial, strong acceptance of evolution denial, strong acceptance of science denial with regard to contraception). We also see vaccine denial and a variety of other science denial movements that are less clearly associated with any one political ideology (the Democratic party and "liberal" media outlets most certainly do not uniformly endorse vaccine denial nor regularly print editorials from vaccine denialists, although, of course, the Huffington Post is well known to do so).
On the other hand, at least we ostensibly do not pay taxes to support purely religious schools that refuse to teach the basics, and that is something we have to keep fighting to maintain.
As for other politics, I will briefly add the following...
I strongly oppose some Israeli actions (I also strongly oppose many US policies) but I also oppose inhumane actions toward or stereotyping of individual Israelis.
I vehemently oppose any suggestion that any current nation state or jurisdiction should be "destroyed" in any way, or that any group of people should be forcibly removed from their homes. I equally condemn both the "destruction of Israel" and the "transfer" of Palestinians as absurdly inhumane ideas which a civilized person would not entertain.
I do NOT advocate "unconditional" support for Israel or any other foreign nation by the US. Nor does any rational person.
The US gives a great deal of aid to Israel and it is in US interests to encourage Israeli policy that meets international standards. However, although the US has the right to determine how much aid it sends (within the limitations of treaty requirements), and to set trade barriers, the US does not have the right to directly dictate domestic policy to any other nation.
"Peace settlements" that require impossibly good behavior from every individual on one side or the other are not merely doomed to fail, but are designed to be doomed to fail. Setting up a "peace settlement" that gives a heckler's veto to any extremist on either side who can manage to blow himself up in public or launch crude rockets is an exercise in futility. Any workable peace settlement is guaranteed to provoke a transient explosion of desperate violence by extremists, who know that if it isn't blocked, it will become the status quo.
I will not be offended if this is sent to the BW, but would prefer that it could be viewed.
John Kwok · 21 June 2010
robert van bakel · 22 June 2010
Haekal was a genious who saw a connection, but over emphasised it. Today we relise the development of embryoes does indeed mirror evolution. It would be interesting if bio-chemists could test the reaction, combinations, and mutations of say, hox-genes to see when deviations become substantial between different species, phyla.
That said, if the Israeli/Palestinian question is ever amicably settled (fat-chance) what on earth will entertain me on evening news? They deserve one another!
Troy Britain · 22 June 2010
Troy Britain · 25 June 2010
Matt Young · 25 June 2010
Paul Strode · 28 June 2010
I agree, and thanks for bringing this to light. I will rewrite this section more clearly if we ever get the chance at a second edition. My intention wasn't to argue that Sedgwick promoted v. Baer over Haeckel, just to establish the timetable for Haeckel's critics.
P.Z. Meyers said it better here:
http://scienceblogs.com/pharyngula/2007/02/wells_and_haeckels_embryos.php
"Sedgwick, for instance, compiled an extensive list of objections to recapitulation as formulated by von Baer and Haeckel, and specifically rejected it as untenable - in 1894."
Paul Strode · 28 June 2010
It's "Myers"
Sorry P.Z.
Mike Elzinga · 5 July 2010
To the asshole spammer from www.b2bsharing.com:
You are going to be reported to the Federal Trade Commission.
That can put you out of business.