The Zonal Practice of Second-year Students in the Biological Faculty of Moscow University

Gametophytes are easily cultured, even with limited facilities. They can he used to illustrate not only the alternation of generations in plant life cycles but also aspects of reproduction and development not easily studied with other plant material. The approach can be adjusted to the needs of biology and botany classes over a wide range of experience from secondary school to university. Information is presented on: the choice of species; the collection, storage and sowing of spores; the composition of the medium; light and temperature requirements; sampling and staining techniques; and methods for determining quantitative parameters of cell and gametophyte development. The basic fern life cycle, and modifications such as apogamy, are outlined. Attention is drawn to the main features of the reproductive biology of ferns and appropriate investigations are suggested. The advantages of gametophytes as experimental material for studying cellular aspects of development in intact green plants are explained. They include striking but consistent responses to a range of cultural conditions and environmental treatments. Suggestions are given for investigations to stimulate further discussion of a wide range of developmental events.     

Selective bird predation on the peppered moth: the last experiment of Michael Majerus

Colour variation in the peppered moth Biston betularia was long accepted to be under strong natural selection. Melanics were believed to be fitter than pale morphs because of lower predation at daytime resting sites on dark, sooty bark. Melanics became common during the industrial revolution, but since 1970 there has been a rapid reversal, assumed to have been caused by predators selecting against melanics resting on today's less sooty bark. Recently, these classical explanations of melanism were attacked, and there has been general scepticism about birds as selective agents. Experiments and observations were accordingly carried out by Michael Majerus to address perceived weaknesses of earlier work. Unfortunately, he did not live to publish the results, which are analysed and presented here by the authors. Majerus released 4864 moths in his six-year experiment, the largest ever attempted for any similar study. There was strong differential bird predation against melanic peppered moths. Daily selection against melanics (s ≈ 0.1) was sufficient in magnitude and direction to explain the recent rapid decline of melanism in post-industrial Britain. These data provide the most direct evidence yet to implicate camouflage and bird predation as the overriding explanation for the rise and fall of melanism in moths.     

Effects of a larval antipredator response and larval diet on adult phenotype in an aposematic ladybird beetle

Many ladybird beetles respond to a potential predation event by `reflex bleeding' or secreting a noxious defensive chemical that is similar to hemolymph. Both adults and larvae show this response. Reflex bleeding is known to reduce predator attack rates and increase prey survival after an attack, especially when reflex bleeding is employed in combination with other cues such as odor and warning coloration. In this experiment, we examined how variability in the number of reflex bleeding events and food quality in the larval stage of the aposematic ladybird beetle Harmonia axyridis affected elytral color, development time, and terminal size in adults. Effects of reflex bleeding were subtle and may have been influenced by diet treatments. Adult color did not differ between bleed treatment groups but beetles that reflex bled tended to take longer to develop and grow to smaller sizes than control group beetles. There were clear and strong effects of larval diet on adult phenotype: an ad libitum pollen diet resulted in paler adult coloration, shorter development time, and larger adult size relative to a limited-availability aphid diet. Our results suggest that the best environment for producing bright-red coloration may not be the best environment for favorable expression of life history characters, especially under stressful conditions. Interactions between different life history stages of H. axyridis are also discussed. Received: 20 April 1997 / Accepted: 30 September 1997     

Weapon (thorn) automimicry and mimicry of aposematic colorful thorns in plants

In order to further characterize the function of coloration in plants as defense against herbivory, two types of thorn mimicry are described: (1) A unique type of weapon (thorn) automimicry (within the same individual) that was previously known only in animals, and (2) mimicry of aposematic colorful thorns, by colorful elongated and pointed plant organs (buds, leaves and fruit) that, despite their appearance, are not sharp. Some thorny plants including dozens of species of Agave, one species of Aloe and a palm species have thorn-like imprints or colorations on their leaves, constituting thorn automimicry by giving the impression of more extensive thorns. The mimicry of aposematic colorful thorns is a typical case of Batesian mimicry, but the thorn automimicry is a special intra-organismic Batesian mimicry. I propose that both types of mimicry serve as anti-herbivore mechanisms.     

Why do some thorny plants resemble green zebras?

The rosette and cauline leaves of the highly thorny winter annual plant species of the Asteraceae in Israel (Silybum marianum) resemble green zebras. The widths of typical variegation bands were measured and found to be highly correlated with leaf length, length of the longest spines at leaf margins and the number of spines along leaf circumference. Thus, there is a significant correlation between the spinyness and strength of variegation. I propose that this is a special case of aposematic (warning) coloration.     

On the selective forces acting in the industrial melanism of Biston and Oligia moths (Lepidoptera: Geometridae and Noctuidae)

Melanic and typical morphs of Biston betularius (L.), Oligia latruncula (D. & S.) and 0. strigilis (L.) made choices between vertical trunks and horizontal branches, sprayed with white and black paints, in a transparent plastic cylinder in natural illumination. The moths settled in exposed positions. In neither Biston nor Oligia did the choice for white/black backgrounds differ between the morphs. Biston moths settle on narrow branches (not on twigs) with the body at right angles to the longitudinal axis of the branch. The Oligias showed an asymmetrical light reaction: one eye is kept in shadow so that they settle as a continuation of an irregularity of the surface, often of a lichen.
In nature, Biston betularius probably rests high up in the canopies, on the under surfaces of horizontal branches. The visual selection acting on the morphs is expected to be less intensive than that measured on tree trunks. The mark-release-recapture results of Kettlewell (1955a, 1956) do not show any qualitative change during the self-determination of the moths but the material is too limited for firm conclusions.
Newly-hatched Biston males take off straight from the trunk where they have expanded their wings but the females may climb higher in the tree. A hypothesis is presented to explain the black-and-white coloration of f. carbonaria : the short-winged moths climbing up the trunks might deter bird predation.     

Retelling Experiments: H.B.D. Kettlewell's Studies of Industrial Melanism in Peppered Moths

H. B. D. Kettlewell's field experiments on industrial melanism in the peppered moth, Biston betularia, have become the best known demonstration of natural selection in action. I argue that textbook accounts routinely portray this research as an example of controlled experimentation, even though this is historically misleading. I examine how idealized accounts of Kettlewell's research have been used by professional biologists and biology teachers. I also respond to some criticisms of David Rudge to my earlier discussions of this case study, and I question Rudge's claims about the importance of purely observational studies for the eventual acceptance and popularization of Kettlewell's explanation for the evolution of industrial melanism.     

Changing views on melanic moths

The rapid rise in frequency of melanic morphs in several moth species, especially the peppered moth Biston betukria , in industrial regions during the 19th century, and the subsequent rapid decline, indicate the action of strong selection. There has recently been a tendency to criticise and question all aspects of research on industrial melanism, including the experiments which suggest that selective predation plays an important part in the changes. These experiments are reexamined, together with evidence for changes in appearance of tree surfaces and for relation of initial melanic frequency to subsequent rate of decline. It is suggested that intense pollution may have been required to drive the carbonaria morph to a high frequency, with frequency patterns over a mosaic environment smoothed by migration. Improvements in these extreme locations then triggered the decline, with litde indication of the environmental changes in areas of moderate pollution. Reasons for criticism of past work are discussed. Industrial melanism continues to provide an exceptional opportunity to analyse a pattern of selection and change in gene frequency.     

Melanic through nature or nurture: genetic polymorphism and phenotypic plasticity in Harmonia axyridis

Abstract Individuals can adapt to heterogeneity in their environment through either local adaptation or phenotypic plasticity. Colour forms of the ladybird Harmonia axyridis are a classic example of local adaptation, in which the frequency of melanic forms varies greatly between populations. In some populations, there are also large seasonal changes in allele frequency, with melanism being costly in summer and beneficial in winter. We report that the non‐melanic morph of H. axyridis dramatically increases its degree of melanization at cold temperatures. Furthermore, there is genetic variation in reaction norms, with different families responding to temperature in different ways. Variation at different spatial and temporal scales appears to have selected for either genetic or phenotypically plastic adaptations, which may be important in thermoregulation. As melanism is known to have a large effect on fitness in H. axyridis, this plasticity of melanization may have hastened its spread as an invasive species.     

Intraspecific variation in wing and pupal melanization in copper butterflies (Lepidoptera: Lycaenidae)

Melanin is a widespread pigment causing variation in skin darkness, with darker phenotypes typically reaching higher equilibrium temperatures than lighter ones. Therefore, selection is expected to favour darker phenotypes in colder environments. In the present study, we show intraspecific variation in pupal (and wing) melanization along an altitudinal gradient in two species of copper butterflies. Both, pupal and wing melanization increased with increasing altitude. Consistent with the thermal melanism hypothesis, darker (high-altitude) pupae reached higher equilibrium temperatures than paler (low-altitude) ones. However, as temperature differences were rather small despite pronounced differences in melanization, we cannot rule out that factors (e.g. ultraviolet protection, disease resistance) other than temperature comprise the principal selective agents. Mechanistically, variation in melanization might be related to variation in hormone titres, as demonstrated by low-altitude pupae showing higher ecdysteroid and juvenile hormone titres compared to high-altitude ones. Furthermore, we report sex differences in wing melanization, with males being darker than females, which is potentially related to a higher flight activity of males.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 301–312.