01.04.2015 Views

The Questions of Developmental Biology

The Questions of Developmental Biology

The Questions of Developmental Biology

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

By this transfer <strong>of</strong> induction from an external inducer to an internal inducer, a trait that<br />

had been induced by the environment became part <strong>of</strong> the genome <strong>of</strong> the organism and could be<br />

selected. Waddington called this phenomenon "genetic assimilation," while Schmalhausen (1949)<br />

called it "stabilizing selection." Both scientists had used orthodox embryology and orthodox<br />

genetics to explain phenomena that had been considered cases <strong>of</strong> Lamarckian "inheritance <strong>of</strong><br />

acquired characteristics."<br />

A shift from environmental stimulus to genetic stimulus might explain sex determination<br />

in Menidia and caste determination in ants. Similarly, the preexisting developmental plasticity <strong>of</strong><br />

arm length in feeding echinoderm larvae may have bridged the transition from pluteus (feeding)<br />

larvae to larvae that lack ciliated arms. <strong>The</strong> change in the allocation <strong>of</strong> resources between larval<br />

and juvenile structures parallels that seen where the food reserves are stored in the egg. Thus, the<br />

changes already present as adaptations to external food resources may have become genetically<br />

fixed in those species whose larvae do not need to hunt their food (Strathmann et al. 1992).<br />

If genetic assimilation is the genetic fixation <strong>of</strong> one <strong>of</strong> the phenotypes that had been<br />

adaptively expressed, then butterflies would be a good place to look for further examples.<br />

Brakefield and colleagues (1996) showed that they could genetically fix the different morphs <strong>of</strong><br />

the adaptive polyphenism <strong>of</strong> Bicyclus, and Shapiro (1976) has shown that the short-day (coldweather)<br />

adaptive phenotype <strong>of</strong> several butterflies is the same as the single genetically produced<br />

phenotype <strong>of</strong> related species or subspecies living at higher altitudes or latitudes.<br />

Genetic assimilation may play an important role in providing a bias for evolutionary<br />

change. If an organism inherits a reaction norm, the developmental pathways leading to a<br />

particular phenotype are already in place, and all that evolution need do is supply a constant<br />

initiator <strong>of</strong> those pathways. In the next chapter, we will discuss some <strong>of</strong> the molecular evidence<br />

for genetic assimilation.<br />

*And until this century, writers were recognized by the calluses on their fingers. (Thus, from observing his fingers,<br />

Sherlock Holmes correctly surmised that the red-headed man had been hired as a scrivener.)<br />

Environmental Disruption <strong>of</strong> Normal Development<br />

From what has been said so far in this chapter, it is clear that the instructions for<br />

development do not reside wholly in the genes or even in the zygote. <strong>The</strong> developing organism is<br />

<strong>of</strong>ten sensitive to cues from the environment. However, this sensitivity makes the organism<br />

vulnerable to environmental changes that can disrupt development.<br />

If you think it is amazing that any<br />

one <strong>of</strong> us survives to be born, you are<br />

correct.It is estimated that one-half to twothirds<br />

<strong>of</strong> all human conceptions do not<br />

develop successfully to term (Figure<br />

21.20). Many <strong>of</strong> these embryos express<br />

their abnormality so early that they fail to<br />

implant in the uterus. Others implant but<br />

fail to establish a successful pregnancy.

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!