Epiallelic variation in Arabidopsis thaliana.

Genotype is the primary determinate of phenotype. During the past two decades, however, there has been an emergent recognition of the epigenotype, a separate layer of heredity distinct from the primary DNA sequence that can have profound effects on phenotype. The epigenotype is a collection of chemical modifications to the DNA and nucleosomes in conjunction with noncoding RNA transcripts, and together these epigenetic marks act as a potent and expansive regulatory system for controlling gene expression. In this review, we discuss our current understanding of variation in epigenotype in the model plant Arabidopsis and how allelic differences attributable to epigenetic changes, or epialleles, can affect phenotype. We discuss examples of epialleles that have been created in the laboratory and others that have been identified in natural populations, because these two models provide complementary information regarding the genetic pathways, mechanisms of transmission, and biological and evolutionary context for the role of the epigenotype in phenotypic variation.

Oral tool use by captive orangutans (Pongo pygmaeus).

Eight captive orangutans (Pongo pygmaeus) were given wooden blocks embedded with raisins and bamboo as raw material for tool making in a study of manual laterality. In about three quarters of the raisin extraction bouts, the orangutans held the tool in the lips or teeth rather than in their hands. Three adult males and 2 adult females showed extreme (> or =92%) preference for oral tool use, a subadult male and an adult female used oral tools about half the time, and 1 adult female preferred manual tool use. Most oral tool users made short tools (approx. 4-10 cm long) that were held in the lips and (probably) supported by the tongue. Preference for oral tool use does not correlate with body weight, age or sex, but it may be related to hand size or individual preference. This is the first report of customary oral tool use as the norm in captive orangutans; it resembles the behavioral patterns reported by van Schaik et al. and Fox et al. in nature.

Expansin message regulation in parasitic angiosperms: marking time in development.

Parasitic strategies are widely distributed across the angiosperms and are estimated to have evolved at least eight different times. Within the obligate hemiparasitic and holoparasitic members, elaborate strategies for host selection have emerged. Here, we demonstrate that in the parasitic Scrophulariceae Striga asiatica, for which signal-mediated host detection is critical, expansin mRNA provides a reliable and accurate downstream molecular marker for the transition to the parasitic mode. Three different expansin genes, saExp1, saExp2, and saExp3, are regulated by xenognostic quinones. saExp3 appears to function as a seedling expansin, and its mRNA is depleted within minutes after induction of the host attachment organ. saExp1 and saExp2 share less homology with the known expansins, and their transcripts accumulate linearly over a critical induction period. The regulation of these genes suggests that the resources for developmental commitment must accumulate to a defined threshold before commitment to organogenesis is terminal. When the induction signal is removed prematurely, the accumulated message decays with a time constant that correlates with the time required for additional signal exposures to reinduce parasitic development. These results suggest that sophisticated controls exist for the accumulation of the necessary components for terminal commitment to the parasitic mode. Furthermore, building on the redox dependence of the inducing signal, they suggest a model akin to a "molecular capacitor" for clocking organogenesis in S. asiatica.

A mechanism for inducing plant development: the genesis of a specific inhibitor.

Parasitic strategies are widely distributed in the plant kingdom and frequently involve coupling parasite organogenesis with cues from the host. In Striga asiatica, for example, the cues that initiate the development of the host attachment organ, the haustorium, originate in the host and trigger the transition from vegetative to parasitic mode in the root meristem. This system therefore offers a unique opportunity to study the signals and mechanisms that control plant cell morphogenesis. Here we establish that the biological activity of structural analogs of the natural inducer displays a marked dependence on redox potential and suggest the existence of a semiquinone intermediate. Building on chemistry that exploits the energetics of such an intermediate, cyclopropyl-p-benzoquinone (CPBQ) is shown to be a specific inhibitor of haustorial development. These data are consistent with a model where haustorial development is initiated by the completion of a redox circuit.