THE ULTRASTRUCTURE OF SCENEDESMUS (CHLOROPHYCEAE). I. SPECIES WITH THE “RETICULATE” OR “WARTY” TYPE OF ORNAMENTAL LAYER1

The ultrastructure of the wall layers and ornamentative features of Scenedesmus pannonicus and S. longus are described using carefully correlated freeze-etched replicas, thin sections and scanning electron micrographs. The two species arc enclosed by different types of ornamented layers, S. pannonicus by the tightly filling, “warty” layer and S. longus by the loosely fitting, “reticulate” layer, held off the coenobium by 2 types of tubular propping spikelets and rosettes. The reticulate layer has an intricate substructure, especially when studied with freeze-etching. Its inner and outer surfaces appear different, as is its attachment to the 2 types of spikelets. Whole cells of S. longus subjected to acetolysis lack the cellulose wall and cytoplasm, but all other surface features survive, including the Trilaminar Sheath (TLS); this ornamentation cannot be “pectic.” The cellulose wall and ornamentation is unaffected by boiling water alone. Boiling in 6n NaOH removes the surface ornamentation, but the TLS and wall remain; the possibility that these features contain silica is discussed. The terminal spines of both species consist of closely packed spikelets enclosed within a skin of hexagonally-packed subunits. Similar subunits are seen in the propping spikelets of S. longus, and in the rows or “combs” of laterally fused spikelets of S. pannonicus. The warty layer of S. pannonicus is tightly appressed to the TLS except close to where the cells are joined, where it is suspended free. It is composed of a layer of globular subunits, and small indentations form the warts. Single, evenly distributed warts characterize the freely suspended sections of the warty layer, and the layer that encloses young coenobia soon after they have been formed: in contrast, the warts are clumped over the surface of older and larger colonies. Some of the single warts form characteristic double rows, but these latter remain single even on older cells. The surface structure of the warty layer, TLS, and plasmalemma are revealed by the freeze-etching process.     

THE CONTRIBUTION OF GENE MOVEMENT TO THE “TWO RULES OF SPECIATION”

The two “rules of speciation”—the Large X‐effect and Haldane's rule—hold throughout the animal kingdom, but the underlying genetic mechanisms that cause them are still unclear. Two predominant explanations—the “dominance theory” and faster male evolution—both have some empirical support, suggesting that the genetic basis of these rules is likely multifarious. We revisit one historical explanation for these rules, based on dysfunctional genetic interactions involving genes recently moved between chromosomes. We suggest that gene movement specifically off or onto the X chromosome is another mechanism that could contribute to the two rules, especially as X chromosome movements can be subject to unique sex‐specific and sex chromosome specific consequences in hybrids. Our hypothesis is supported by patterns emerging from comparative genomic data, including a strong bias in interchromosomal gene movements involving the X and an overrepresentation of male reproductive functions among chromosomally relocated genes. In addition, our model indicates that the contribution of gene movement to the two rules in any specific group will depend upon key developmental and reproductive parameters that are taxon specific. We provide several testable predictions that can be used to assess the importance of gene movement as a contributor to these rules in the future.     

COMPETITION BETWEEN SEGREGATION DISTORTERS: COEXISTENCE OF “SUPERIOR” AND “INFERIOR” HAPLOTYPES AT THE t COMPLEX

By means of population genetical models, we investigate the competition between sex-specific segregation distorters. Although the models are quite general, they are motivated by a specific example, the t complex of the house mouse. Some variants at this gene complex, the t haplotypes, distort Mendelian segregation in heterozygous males in their favor. The selective advantage at the gamete level is counterbalanced by strong negative fitness effects at the individual level (male sterility or even lethality in both sexes). A plethora of different t haplotypes has been found, both in the field and in the lab. Up to now, however, models have focused on the equilibrium frequency of a single t haplotype. In contrast, we explicitly model the competition between several t haplotypes. A deterministic model for a large, well-mixed population predicts a surprisingly high degree of polymorphism. Haplotypes with seemingly inferior fitness characteristics may easily coexist with “superior” haplotypes. For instance, a lethal haplotype with a low segregation ratio may stably coexist with a sterile haplotype with a high segregation ratio. Stable coexistence is even possible for haplotypes with a segregation disadvantage. A simple stochastic model shows that the same principles apply in the context of a structured metapopulation. Although counterintuitive at first sight, all our results can be explained by the fact that segregation distorters have an inherent advantage when they are rare. We conclude that fitness comparisons are not sufficient to predict the outcome of competition when selective forces are acting at different levels.     

THE EVOLUTION OF PHENOTYPIC CORRELATIONS AND “DEVELOPMENTAL MEMORY”

Development introduces structured correlations among traits that may constrain or bias the distribution of phenotypes produced. Moreover, when suitable heritable variation exists, natural selection may alter such constraints and correlations, affecting the phenotypic variation available to subsequent selection. However, exactly how the distribution of phenotypes produced by complex developmental systems can be shaped by past selective environments is poorly understood. Here we investigate the evolution of a network of recurrent nonlinear ontogenetic interactions, such as a gene regulation network, in various selective scenarios. We find that evolved networks of this type can exhibit several phenomena that are familiar in cognitive learning systems. These include formation of a distributed associative memory that can “store” and “recall” multiple phenotypes that have been selected in the past, recreate complete adult phenotypic patterns accurately from partial or corrupted embryonic phenotypes, and “generalize” (by exploiting evolved developmental modules) to produce new combinations of phenotypic features. We show that these surprising behaviors follow from an equivalence between the action of natural selection on phenotypic correlations and associative learning, well‐understood in the context of neural networks. This helps to explain how development facilitates the evolution of high‐fitness phenotypes and how this ability changes over evolutionary time.     

THE “BIOTYPES” OF THE RICE BROWN PLANTHOPPER,NILAPARVATA LUGENS

From previous studies, the biological nature of the so-called biotypes of N. lugens is obscure. Experiments on biotypes 1, 2 and 3 from the Philippines demonstrated that they lacked significant breeding barriers. Inbred biotype cultures showed considerable variation and overlap between each other in virulence. A field population from the Philippines was similarly variable and included individuals which could be attributed to different biotypes. It is concluded that the use of the term biotype in this species is biologically misleading and undesirable.

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Résumé La nature biologique de ce qui a été appelé des biotypes de Nilaparvata lugens, paraît obscure si l'on se réfère aux études antérieures. Des expériences sur les biotypes 1, 2 et 3 des Philippines révèlent qu'ils ne présentent pas de barrières sexuelles significatives. Les élevages consanguins de biotypes présentent une variabilité considérable, et les virulances des biotypes se chevauchent les unes les autres. De même une population récoltée dans les çhamps aux Philippines était hétérogène et comprenait des individus qui auraient pu être rattachés à différents biotypes. On peut en conclure que le terme de biotype est, pour cette espèce, inapproprié et indésirable.

     

THE GENETICS OF THE “BIOTYPES” OF THE RICE BROWN PLANTHOPPER,NILAPARVATA LUGENS

As the resistance of rice varieties to the brown planthopper is based on major genes it has been widely assumed that there is a gene for gene correspondence between resistance on the part of the plant and virulence on the part of the hopper. However, the mode of inheritance and response of the biotypes to selection, together with the previously reported wide variation within each biotype and large overlap between them in virulence, is all consistent with polygenic determination of virulence.

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Résumé Comme la résistance des variétés de riz à N. lugens est due à des gènes majeurs, on considère généralement qu'il y a une correspondance gène à gène entre la résistance de la plante et la virulence de l'insête. Cependant, le type d'héritabilité et la réponse des biotypes à la sélection, ainsi que la grande variabilité antérieurement signalée dans chaque biotype et l'important chevauchement de leur virulence, sont tous conformes à une détermination polygénique de la virulence.

     

THE EFFECT OF CORPORA ALLATA ON SEXUAL BEHAVIOUR AND “ADULT DIAPAUSE” IN MALES OF THE RED LOCUST

Males of the red locust, Nomadacris septemfasciata, allatectomized 3–11 days after the moult to the adult stage did not exhibit any sexual behaviour and remained in a state comparable to adult diapause, while all the sham-allatectomized controls exhibited distinct sexual behaviour. Implantations of four pairs of active corpora allata (originating from sexually mature males of Locusta migratoria migratorioides) into the allatectomized males of the red locust led to sexual behaviour; in contrast, all those allatectomized males in which shamimplantations (appropriate surgical treatment without actual implantation of corpora allata) were performed remained in adult diapause showing no sexual behaviour.Comparison of the results with other data revealed that in some species of Acridids the corpora allata completely control (in a direct or indirect manner) male sexual behaviour, while in other species of the same family this control is either incomplete or perhaps does not exist at all. The presence or absence of a complete endocrine control of male sexual behaviour may perhaps be correlated with the fact that the species involved has or has not the ability to undergo a state of adult diapause.

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Zusammenfassung Männchen der Roten Heuschrecke, Nomadacris septemfasciata, die 3–11 Tage nach der Häutung zur Imago allatektomiert worden waren, zeigten keinerlei sexuelle Aktivität und verharrten in einem der Adult-Diapause vergleichbaren Zustand, während alle scheinallatektomierten Kontrolltiere ausgeprägtes Sexualverhalten aufwiesen. Implantationen von 4 Paar Corpora allata (aus geschlechtsreifen Männchen von Locusta migratoria migratorioides stammend) in allatektomierte Männchen der Roten Heuschrecke führten zu Sexualverhalten; im Gegensatz dazu verharrten alle diejenigen allatektomierten Männchen, bei welchen Schein-Implantationen durchgeführt wurden (entsprechende operative Behandlung ohne Implantation von Corpora allata) in Adult-Diapause und zeigten keine sexuelle Aktivität.Vergleich der Ergebnisse mit anderen Daten lassen erkennen, daß bei einigen Acridier-Arten die Corpora allata das männliche Sexualverhalten (auf direkte oder indirekte Weise) beeinflussen, während dieser Einfluß bei anderen Arten der gleichen Familie entweder unvollständig ist oder überhaupt nicht auftritt. Vorkommen oder Fehlen einer vollständigen endokrinen Kontrolle des männlichen Sexualverhaltens mag vielleicht von dem Umstand abhängen, ob die betreffende Art die Fähigkeit hat, ein Stadium adulter Diapause zu durchlaufen oder nicht.

     

THE ROLE OF PHYLLOTACTIC PATTERN AS A “DEVELOPMENTAL CONSTRAINT” ON THE INTERCEPTION OF LIGHT BY LEAF SURFACES

Computer simulations of model plants are used to assess the influence of leaf shape, size, and pattern of arrangement (= phyllotaxy) on the direct solar radiation intercepted by leaf surfaces. Changes in phyllotaxy significantly influence light interception (and, by inference, net assimilation rate) for rosette growth habits. However, changes in leaf shape and orientation and in stem length can compensate for the negative effects of leaf overlap produced by phyllotactic patterns. Phyllotaxy is viewed as a developmental limiting factor in photobiology that may necessitate compensatory changes in other morphological features not directly controlled by patterns of leaf initiation. This distinguishes it from functioning as a “developmental constraint” sensu stricto and may provide a paradigm for other features in plant evolution.     

THE EVOLUTION OF COSTLY MATE PREFERENCES II. THE “HANDICAP” PRINCIPLE

We use a general additive quantitative genetic model to study the evolution of costly female mate choice by the “handicap” principle. Two necessary conditions must be satisfied for costly preference to evolve. The conditions are (i) biased mutation pressure on viability and (ii) a direct relationship between the degree of expression of the male mating character and viability. These two conditions explain the success and failure of previous models of the “handicap” principle. Our model also applies to other sources of fitness variation like migration and host-parasite coevolution, which cause effects equivalent to biased mutation.