Functional anatomy of the ovule in Genlisea with remarks on ovule evolution in Lentibulariaceae

The Lentibulariaceae are highly evolved and specialized carnivorous angiosperms displaying not only unusual morphology and embryology but also specific changes in the genome and chromosomes as large as bacterial chromosomes. Comparative study of the morphology and detailed anatomy of the ovule in the genera Genlisea, Utricularia, and Pinguicula should shed new light on the phylogeny of this family. The clade Genlisea + Utricularia is sister to the genus Pinguicula, which is considered the most primitive taxon within Lentibulariaceae. Thus we should expect the ovules of Genlisea to be more similar to those of the more closely related genus Utricularia than to Pinguicula. Surprisingly, the ovules of Genlisea retain characters (free funiculus, ES remaining in the ovule) in common with Pinguicula, presumably inherited from a common ancestor. Genlisea ovules have only one main character in common with subgenus Polypompholyx (Utricularia): a well-developed funiculus. There are differences between the ovules of the subgenera Genlisea and Tayloria. In subgenus Genlisea the micropyle tends to be closer to the funiculus and the ovule forms an unusual jacket-like nutritive tissue of integumental origin. The most specialized ovules in Lentibulariaceae evolved in the genus Utricularia. The special chalazal nutritive tissue in Genlisea and Utricularia is simply a hypostase.     

The origin of the ovule

Of several theories for the origin of the ovule advanced in this century and based largely on fossil evidence, the telomic concept is widely considered the most plausible. Its principal tenet is the evolution of the integument through fusion of sterile branches or telomes around a terminal megasporangium. The only point of agreement in these theories is that the entire nucellus is a megasporangium that retains a single megaspore and the endosporic female gametophyte. Their differences center on the origin of the integument. A new concept offered here on evidence from ovules of both Paleozoic and extant seed plants significantly alters the telomic theory. It proposes that the nucellus is a sporangiophore of stem origin that bears a terminal megasporangium; that at least some of the fused integumentary telomes were fertile; and that among all features cited to characterize ovules, the unique nature of the retained megaspore alone defines the seed habit. Changes in the seed plant megaspore that extended the period of nutrient absorption over the whole course of female gametophyte development, along with complex physiological changes in the nucellus, were probably achieved along a single phylogenetic line beginning in a Late Devonian population of progymnosperms. For such a combination of events to have occurred more than once is highly unlikely, and, therefore, a monophyletic origin for seed plants is proposed. Several primitive features in ovule structure, some not evidenced since the Lower Carboniferous Period, occur in a mutant form of Arabidopsis thaliana isolated from genetically transformed plants. Their recurrence provides additional support for the proposed concept of ovule origin and also suggests that the genetic mechanisms for expression of primitive features in advanced taxa could be initiated in each case by mutation of a single homeotic gene.     

Interaction between octopamine and proctolin on the oviducts of Locusta migratoria

The biogenic amine octopamine and the pentapeptide proctolin are two important neuroactive chemicals that control contraction of the oviducts of the African locust Locusta migratoria. The physiological responses and signal transduction pathways used by octopamine and proctolin have been well characterized in the locust oviducts and this therefore provides the opportunity to examine the interaction between these two pathways. Octopamine, via the intracellular messenger adenosine 3',5'-cyclic monophosphate (cyclic AMP), inhibits contraction of the oviducts, while proctolin, via the phosphoinositol pathway, stimulates contraction. We have examined the physiological response of the oviducts to combinations of octopamine and proctolin and also looked at how combinations of these affect one of the main intracellular mediators of the octopamine response, namely cyclic AMP. It was found that application of octopamine to the oviducts led to a dose-dependent reduction in tonus of the muscle and also a decrease in the amplitude and frequency of spontaneous phasic contractions. Octopamine-induced relaxation was enhanced in the presence of the phosphodiesterase inhibitor, 3-isobutyl-1-methylxanthine (IBMX). Octopamine was also able to inhibit proctolin-induced contractions of the oviducts in a dose-dependent manner. A 10(-9) M proctolin-induced contraction was inhibited by 83% in the presence of 10(-5) M octopamine, and was completely inhibited in the presence of 10(-5) M octopamine plus 5x10(-4) M IBMX. Octopamine led to a dose-dependent increase in cyclic AMP content as measured by radioimmunoassay. In the presence of 10(-9) M proctolin, this octopamine-induced increase in cyclic AMP was reduced by as much as 60%. Proctolin also caused a dose-dependent decrease in the cyclic AMP elevation produced by 5x10(-6) M octopamine. These results indicate that octopamine and proctolin can antagonize each other's physiological response when added in combination, and that proctolin is able to modulate the response of the oviducts to octopamine by influencing cyclic AMP levels.