Sex Chromosome Evolution The fly genus Drosophilia has repeatedly generated evolutionarily new sex chromosomes. To understand the changes shaping the X and Y chromosomes, Zhou and Bachtrog (p. 341), sequenced the genome of D. miranda, which formed neo-X and neo-Y chromosomes approximately 1 million years ago. The data illuminate the ongoing conflict between selection for male and female function on the sex chromosomes and show that Y chromosome evolution is characterized both by a loss of gene function and selection for male-specific adaptations in genes beneficial to male functions. Evolutionarily new X and Y chromosomes evidence how selection for sexual function shapes sex chromosomes. Most species’ sex chromosomes are derived from ancient autosomes and show few signatures of their origins. We studied the sex chromosomes of Drosophila miranda, where a neo-Y chromosome originated only approximately 1 million years ago. Whole-genome and transcriptome analysis reveals massive degeneration of the neo-Y, that male-beneficial genes on the neo-Y are more likely to undergo accelerated protein evolution, and that neo-Y genes evolve biased expression toward male-specific tissues—the shrinking gene content of the neo-Y becomes masculinized. In contrast, although older X chromosomes show a paucity of genes expressed in male tissues, neo-X genes highly expressed in male-specific tissues undergo increased rates of protein evolution if haploid in males. Thus, the response to sex-specific selection can shift at different stages of X differentiation, resulting in masculinization or demasculinization of the X-chromosomal gene content.

Sex-Specific Adaptation Drives Early Sex Chromosome Evolution In Drosophila

Genomic analysis of many nonmodel species has uncovered an incredible diversity of sex chromosome systems, making it possible to empirically test the rich body of evolutionary theory that describes each stage of sex chromosome evolution. Classic theory predicts that sex chromosomes originate from a pair of homologous autosomes and recombination between them is suppressed via inversions to resolve sexual conflict. The resulting degradation of the Y chromosome gene content creates the need for dosage compensation in the heterogametic sex. Sex chromosome theory also implies a linear process, starting from sex chromosome origin and progressing to heteromorphism. Despite many convergent genomic patterns exhibited by independently evolved sex chromosome systems, and many case studies supporting these theoretical predictions, emerging data provide numerous interesting exceptions to these long-standing theories, and suggest that the remarkable diversity of sex chromosomes is matched by a similar diversity in their evolution. For example, it is clear that sex chromosome pairs are not always derived from homologous autosomes. In addition, both the cause and the mechanism of recombination suppression between sex chromosome pairs remain unclear, and it may be that the spread of recombination suppression is a more gradual process than previously thought. It is also clear that dosage compensation can be achieved in many ways, and displays a range of efficacy in different systems. Finally, the remarkable turnover of sex chromosomes in many systems, as well as variation in the rate of sex chromosome divergence, suggest that assumptions about the inevitable linearity of sex chromosome evolution are not always empirically supported, and the drivers of the birth-death cycle of sex chromosome evolution remain to be elucidated. Here, we concentrate on how the diversity in sex chromosomes across taxa highlights an equal diversity in each stage of sex chromosome evolution.

Sex Chromosome Evolution: So Many Exceptions to the Rules.

Butterfly eyespot colour patterns are a key example of how a novel trait can appear in association with the co-option of developmental patterning genes. Little is known, however, about how, or even whether, co-opted genes function in eyespot development. Here we use CRISPR/Cas9 genome editing to determine the roles of two co-opted transcription factors that are expressed during early eyespot determination. We found that deletions in a single gene, spalt, are sufficient to reduce or completely delete eyespot colour patterns, thus demonstrating a positive regulatory role for this gene in eyespot determination. Conversely, and contrary to previous predictions, deletions in Distal-less (Dll) result in an increase in the size and number of eyespots, illustrating a repressive role for this gene in eyespot development. Altogether our results show that the presence, absence and shape of butterfly eyespots can be controlled by the activity of two co-opted transcription factors.

/pdf/genome-editing-in-butterflies-reveals-that-spalt-promotes-1p9tv5q76z.pdf

Genome editing in butterflies reveals that spalt promotes and Distal-less represses eyespot colour patterns

Eyespots on the wings of nymphalid butterflies represent colorful examples of pattern formation, yet the developmental origins and mechanisms underlying eyespot center differentiation are still poorly understood. Using CRISPR-Cas9 we re-examine the function of Distal-less (Dll) as an activator or repressor of eyespots, a topic that remains controversial. We show that the phenotypic outcome of CRISPR mutations depends upon which specific exon is targeted. In Bicyclus anynana, exon 2 mutations are associated with both missing and ectopic eyespots, and also exon skipping. Exon 3 mutations, which do not lead to exon skipping, produce only null phenotypes, including missing eyespots, lighter wing coloration and loss of scales. Reaction-diffusion modeling of Dll function, using Wnt and Dpp as candidate morphogens, accurately replicates these complex crispant phenotypes. These results provide new insight into the function of Dll as a potential activator of eyespot development, scale growth and melanization, and suggest that the tuning of Dll expression levels can generate a diversity of eyespot phenotypes, including their appearance on the wing.This article has an associated 'The people behind the papers' interview.

/pdf/activation-of-butterfly-eyespots-by-distal-less-is-n77kikwr8g.pdf

Activation of butterfly eyespots by Distal-less is consistent with a reaction-diffusion process.

Wingless is a positive regulator of eyespot color patterns in Bicyclus anynana butterflies.

The border ocellus, or eyespot, is a conspicuous color pattern element in butterfly wings. For two decades, it has been hypothesized that transcription factors such as Distal-less (Dll) are responsible for eyespot pattern development in butterfly wings, based on their expression in the prospective eyespots. In particular, it has been suggested that Dll is a determinant for eyespot size. However, functional evidence for this hypothesis has remained incomplete, due to technical difficulties. Here, we show that ectopically expressed Dll induces ectopic elemental color patterns in the adult wings of the blue pansy butterfly, Junonia orithya (Lepidoptera, Nymphalidae). Using baculovirus-mediated gene transfer, we misexpressed Dll protein fused with green fluorescent protein (GFP) in pupal wings, resulting in ectopic color patterns, but not the formation of intact eyespots. Induced changes included clusters of black and orange scales (a basic feature of eyespot patterns), black and gray scales, and inhibition of cover scale development. In contrast, ectopic expression of GFP alone did not induce any color pattern changes using the same baculovirus-mediated gene transfer system. These results suggest that Dll plays an instructive role in the development of color pattern elements in butterfly wings, although Dll alone may not be sufficient to induce a complete eyespot. This study thus experimentally supports the hypothesis of Dll function in eyespot development.

/pdf/distal-less-induces-elemental-color-patterns-in-junonia-3q8yzvstc9.pdf

Distal-less induces elemental color patterns in Junonia butterfly wings.

Sex chromosomes and sex determining genes can evolve fast, with the sex-linked chromosomes often differing between closely related species. Population genetics theory has been developed and tested to explain the rapid evolution of sex chromosomes and sex determination. However, we do not know why the sex chromosomes are divergent in some taxa and conserved in others. Addressing this question requires comparing closely related taxa with conserved and divergent sex chromosomes to identify biological features that could explain these differences. Cytological karyotypes suggest that muscid flies (e.g., house fly) and blow flies are such a taxonomic pair. The sex chromosomes appear to differ across muscid species, whereas they are conserved across blow flies. Despite the cytological evidence, we do not know the extent to which muscid sex chromosomes are independently derived along different evolutionary lineages. To address that question, we used genomic and transcriptomic sequence data to identify young sex chromosomes in two closely related muscid species, horn fly (Haematobia irritans) and stable fly (Stomoxys calcitrans). We provide evidence that the nascent sex chromosomes of horn fly and stable fly were derived independently from each other and from the young sex chromosomes of the closely related house fly (Musca domestica). We present three different scenarios that could have given rise to the sex chromosomes of horn fly and stable fly, and we describe how the scenarios could be distinguished. Distinguishing between these scenarios in future work could identify features of muscid genomes that promote sex chromosome divergence.

/pdf/sex-chromosome-evolution-in-muscid-flies-56orcvet9s.pdf

Sex Chromosome Evolution in Muscid Flies.

It is often desirable but difficult to retrieve information on the mature phenotype of an immature tissue sample that has been subjected to gene expression analysis. This problem cannot be ignored when individual variation within a species is large. To circumvent this problem in the butterfly wing system, we developed a new surgical method for removing a single forewing from a pupa using Junonia orithya; the operated pupa was left to develop to an adult without eclosion. The removed right forewing was subjected to gene expression analysis, whereas the non-removed left forewing was examined for color patterns. As a test case, we focused on Distal-less (Dll), which likely plays an active role in inducing elemental patterns, including eyespots. The Dll expression level in forewings was paired with eyespot size data from the same individual. One third of the operated pupae survived and developed wing color patterns. Dll expression levels were significantly higher in males than in females, although male eyespots were smaller in size than female eyespots. Eyespot size data showed weak but significant correlations with the Dll expression level in females. These results demonstrate that a single-wing removal method was successfully applied to the butterfly wing system and suggest the weak and non-exclusive contribution of Dll to eyespot size determination in this butterfly. Our novel methodology for establishing correspondence between gene expression and phenotype can be applied to other candidate genes for color pattern development in butterflies. Conceptually similar methods may also be applicable in other developmental systems.

https://bioone.org/journals/zoological-science/volume-33/issue-1/zs150113/A-Single-Wing-Removal-Method-to-Assess-Correspondence-Between-Gene/10.2108/zs150113.pdf

A Single-Wing Removal Method to Assess Correspondence Between Gene Expression and Phenotype in Butterflies: The Case of Distal-less

In this paper, we study the Krylov complexity ($K$) from the planar/inflationary patch of the de Sitter space using the two mode squeezed state formalism in the presence of an effective field having sound speed $c_s$. From our analysis, we obtain the explicit behavior of Krylov complexity ($K$) and lancoz coefficients ($b_n$) with respect to the conformal time scale and scale factor in the presence of effective sound speed $c_s$. Since lancoz coefficients ($b_n$) grow linearly with integer $n$, this suggests that universe acts like a chaotic system during this period. We also obtain the corresponding Lyapunov exponent $\lambda$ in presence of effective sound speed $c_s$. We show that the Krylov complexity ($K$) for this system is equal to average particle numbers suggesting it's relation to the volume. Finally, we give a comparison of Krylov complexity ($K$) with entanglement entropy (Von-Neumann) where we found that there is a large difference between Krylov complexity ($K$) and entanglement entropy for large values of squeezing amplitude. This suggests that Krylov complexity ($K$) can be a significant probe for studying the dynamics of the cosmological system even after the saturation of entanglement entropy.

/pdf/cosmological-krylov-complexity-18m3hyav.pdf

Cosmological Krylov Complexity

In this paper, we study the Krylov complexity in quantum ﬁeld theory and make a connection with the holographic "Complexity equals Volume" conjecture. When Krylov basis matches with Fock basis, for several interesting settings, we observe that the Krylov complexity equals the average particle number showing that complexity scales with volume. Using similar formalism, we compute the Krylov complexity for free scalar ﬁeld theory and ﬁnd surprising similarities with holography. We also extend this framework for ﬁeld theory where an inverted oscillator appears naturally and explore its chaotic behavior.

Kiran Adhikari

Papers

Distal-less induces elemental color patterns in Junonia butterfly wings.

Sex Chromosome Evolution in Muscid Flies.

A Single-Wing Removal Method to Assess Correspondence Between Gene Expression and Phenotype in Butterflies: The Case of Distal-less

Cosmological Krylov Complexity

K rylov C omplexity in Q uantum F ield T heory