Tetrachromacy in a butterfly that has eight varieties of spectral receptors

This paper presents the first evidence of tetrachromacy among invertebrates. The Japanese yellow swallowtail butterfly, Papilio xuthus, uses colour vision when foraging. The retina of Papilio is furnished with eight varieties of spectral receptors of six classes that are the ultraviolet (UV), violet, blue (narrow-band and wide-band), green (single-peaked and double-peaked), red and broad-band classes. We investigated whether all of the spectral receptors are involved in colour vision by measuring the wavelength discrimination ability of foraging Papilio. We trained Papilio to take nectar while seeing a certain monochromatic light. We then let the trained Papilio choose between two lights of different wavelengths and determined the minimum discriminable wavelength difference Deltalambda. The Deltalambda function of Papilio has three minima at approximately 430, 480 and 560nm, where the Deltalambda values approximately 1nm. This is the smallest value found for wavelength discrimination so far, including that of humans. The profile of the Deltalambda function of Papilio can be best reproduced by postulating that the UV, blue (narrow-band and wide-band), green (double-peaked) and red classes are involved in foraging. Papilio colour vision is therefore tetrachromatic.     

Coloured oil droplets enhance colour discrimination

The eyes of most diurnal reptiles and birds contain coloured retinal filters-oil droplets. Although these filters are widespread, their adaptive advantage remains uncertain. To understand why coloured oil droplets appeared and were retained during evolution, I consider both the benefits and the costs of light filtering in the retina. Oil droplets decrease cone quantum catch and reduce the overlap in sensitivity between spectrally adjacent cones. The reduction of spectral overlap increases the volume occupied by object colours in a cone space, whereas the decrease in quantum catch increases noise, and thus reduces the discriminability of similar colours. The trade-off between these two effects determines the total benefit of oil droplets. Calculations show that coloured oil droplets increase the number of object colours that can be discriminated, and thus are beneficial for colour vision.     

Photoreceptor spectral sensitivities in terrestrial animals: adaptations for luminance and colour vision

This review outlines how eyes of terrestrial vertebrates and insects meet the competing requirements of coding both spatial and spectral information. There is no unique solution to this problem. Thus, mammals and honeybees use their long-wavelength receptors for both achromatic (luminance) and colour vision, whereas flies and birds probably use separate sets of photoreceptors for the two purposes. In particular, we look at spectral tuning and diversification among 'long-wavelength' receptors (sensitivity maxima at greater than 500 nm), which play a primary role in luminance vision. Data on spectral sensitivities and phylogeny of visual photopigments can be incorporated into theoretical models to suggest how eyes are adapted to coding natural stimuli. Models indicate, for example, that animal colour vision--involving five or fewer broadly tuned receptors--is well matched to most natural spectra. We can also predict that the particular objects of interest and signal-to-noise ratios will affect the optimal eye design. Nonetheless, it remains difficult to account for the adaptive significance of features such as co-expression of photopigments in single receptors, variation in spectral sensitivities of mammalian L-cone pigments and the diversification of long-wavelength receptors that has occurred in several terrestrial lineages.     

Modelling oil droplet absorption spectra and spectral sensitivities of bird cone photoreceptors

Birds have four spectrally distinct types of single cones that they use for colour vision. It is often desirable to be able to model the spectral sensitivities of the different cone types, which vary considerably between species. However, although there are several mathematical models available for describing the spectral absorption of visual pigments, there is no model describing the spectral absorption of the coloured oil droplets found in three of the four single cone types. In this paper, we describe such a model and illustrate its use in estimating the spectral sensitivities of single cones. Furthermore, we show that the spectral locations of the wavelengths of maximum absorbance (_max) of the short- (SWS), medium- (MWS) and long- (LWS) wavelength-sensitive visual pigments and the cut-off wavelengths (_cut) of their respective C-, Y- and R-type oil droplets can be predicted from the _max of the ultraviolet- (UVS)/violet- (VS) sensitive visual pigment.     

Discrimination of coloured stimuli by honeybees: alternative use of achromatic and chromatic signals

Using a Y-maze experimental set-up, honeybees Apis mellifera were trained to a coloured disc presented against an achromatic background. In subsequent tests they were given a choice between the trained disc and an alternative disc that differed either in its chromatic properties, or in the amount of achromatic green contrast that it produced against the background. Tests were conducted in two experimental situations: one in which discs subtended a visual angle of 30° (as viewed by the bee at the decision point in the Y-maze), and another in which the angle was 6.5° or 5° (depending on the experiment). At the visual angle of 30°, the bees' choice behaviour was governed by the differences in chromatic properties, and not by the differences in the amount of green contrast. With the 6.5°- and 5°-discs, on the other hand, it was governed by the differences in the amount of green contrast, and not by the differences in chromatic properties. Consequently, in the present discrimination task, bees use either chromatic or achromatic cues, depending on the visual angle subtended by the stimuli at the eye. Results of a further experiment, in which the trained disc was tested against discs that produced various amounts of green contrast, confirm the above conclusion and show, in addition, that bees learn the green-contrast difference between a trained and a non-rewarded alternative. Accepted: 25 October 1996     

Biological properties of some nitrogen-fixing associative enterobacteria

Enterobacteria isolates from potato tubers were able to fix nitrogen, to protect plants against phytopathogens and to produce phytohormones thus increasing the plant yield. These isolates were previously phenotypically identified as Erwinia carotovora; however, they differed from typical E. carotovora in a number of biological characteristics and were found to be nonphytopathogenic (avirulent) due to the lack of pectate lyase activity. A data matrix, containing 31 strains and 105 biological characteristics was used for computer cluster analysis. The avirulent strains formed a separate cluster more closely related to Klebsiella spp. strains (with a 0.67 level of similarity) than to typical phytopathogenic bacteria of the E. carotovora group (with a 0.48 level of similarity). A phylogenetic analysis based on restriction polymorphisms of an amplified ribosomal DNA spacer region revealed that the avirulent strains studied here were different from all Erwinia, Klebsiella and other enterobacteria species strains. The AP PCR/hybridization technique showed cross homology of amplified DNA of these avirulent strains and a lack of such homology with the DNA from strains of other species. Numerical taxonomy data, rDNA analysis and AP PCR/hybridization assays confirmed that these avirulent bacteria may be regarded as an independent group of enterobacteria.     

The angular range of achromatic target detection by honey bees

Honeybees Apis mellifera were trained to enter a Y-maze and choose the arm with a rewarded disc presented against a grey background. The alternative arm displayed the unrewarded grey background alone. Training and testing were performed with the rewarding disc subtending different visual angles. The training disc was either achromatic and provided green contrast, or chromatic and provided the same amount of green contrast as the achromatic one. The bee-achromatic disc could be learned and detected by the bees whenever it subtended 5° or 10°, but not if it subtended 30°. The chromatic disc was learned well and detected at all three visual angles. However, at 5° the maximum level of correct choices was ca. 75% with the achromatic disc whilst it was ca. 90% with the chromatic one. Thus, the presence of chromatic contrast enhances considerably the level of correct choices for the same amount of green contrast. The lower threshold of achromatic target detection lies between 3.7° and 5°; the upper threshold between 15° and 10°. At the upper threshold, detection switches from chromatic-based to achromatic-based. Thus, in the context of target detection, the achromatic green contrast channel specialises in the detection of objects of reduced angular size, whilst the chromatic channels are specialised for objects of large angular size. We suggest that achromatic detectors with a centre-surround organisation are involved in the task of detecting achromatic targets. Accepted: 23 February 1998     

Biochemical observation of the rapid mobility of nuclear HMGB1

Formaldehyde-crosslinked and sonicated chromatin fragments were obtained from 15-day chicken embryo erythrocytes and purified on caesium chloride gradients. Polyclonal antibodies raised against chicken HMGB1 were used to immuno-precipitate fragments carrying HMGB1 in two protocols: (1) affinity purified antibodies covalently coupled to agarose beads and (2) diluted antiserum. The DNA of the antibody-bound chromatin was quantified and its sequence content assessed by quantitative real-time PCR to give values of the absolute enrichments generated. Amplicons were monitored within the active beta-globin locus, in the adjacent heterochromatin, in the lysozyme locus (containing an active housekeeping gene and the inactive lysozyme gene) and at the promoter of the inactive ovalbumin gene. For all amplicons the Bound/Input ratio was close to unity, implying no preferential location of HMGB1 on the chromatin. This initially unexpected result can now be understood in the light of the exceptional mobility of HMGB1 revealed by FLIP experiments showing that only 1-2 s are needed for HMGB1 to cross the nucleus: crosslinking times of 1 min were used in the present experiments.     

Hypermethylation of 5′-Region of the Human Calcitonin Gene in Leukemias: Structural Features and Diagnostic Significance

Methylation of the 5-region of the calcitonin gene was investigated in bone marrow and peripheral blood cells of 27 healthy volunteers and 25 leukemic patients. In all patients suffering from various forms of myeloid and lymphoid leukemia, hypermethylation of CpG sequences was observed in this region of the calcitonin gene. Cytosine hypermethylation in the CpG sequence did not involve cytosines of adjacent CpNpG sequences (where N is any nucleoside). The 5-region of the calcitonin gene lacked CpNpG methylation both in healthy controls and in leukemic patients; this apparently represents specific non-alternative type of CpG methylation in the extended DNA sequence. Methylation of the calcitonin gene was monitored in 18 leukemic patients during malignant progression and medical treatment. Hypermethylation of the calcitonin gene was not observed on long-term clinical hematological remission. In ten patients characterized by unstable (or incomplete) remission hypermethylation of the calcitonin gene persisted through the whole period of observation. In relapses, hypermethylation of the calcitonin gene appeared again and in six patients, this molecular relapse being registered 1-8 months before onset of clinical and laboratory signs of disease progression. The leukemia-specific hypermethylation of CpG sequences of the 5-region of the calcitonin gene is a promising prognostic and diagnostic marker of leukemias and might be useful for monitoring of this disease.