This is the oldest English language journal in genetics. Founded by W. Bateson and R. C. Punnett in 1910, the Journal of Genetics was later edited by J. B. S. Haldane; later, Haldane and his wife, Helen Spurway brought the journal with them to India. After his death in 1964, Mrs. Haldane continued publication of the journal, with Madhav Gadgil and H. Sharat Chandra (Indian Institute of Science, Bangalore) and Suresh Jayakar (Laboratorio de Genetica Biochimica ed Evoluzionistica, Pavia) as editors, until her own death in 1977. In 1985, with the permission of Haldane’s heirs, the Indian Academy of Sciences resumed publication of the journal. The journal retains its traditional interest in evolutionary research that is of relevance to geneticists, even if not explicitly genetic in nature.

The journal of genetics

Sex chromosomes evolved from autosomes many times across the eukaryote phylogeny. Several models have been proposed to explain this transition, some involving male and female sterility mutations linked in a region of suppressed recombination between X and Y (or Z/W, U/V) chromosomes. Comparative and experimental analysis of a reference genome assembly for a double haploid YY male garden asparagus (Asparagus officinalis L.) individual implicates separate but linked genes as responsible for sex determination. Dioecy has evolved recently within Asparagus and sex chromosomes are cytogenetically identical with the Y, harboring a megabase segment that is missing from the X. We show that deletion of this entire region results in a male-to-female conversion, whereas loss of a single suppressor of female development drives male-to-hermaphrodite conversion. A single copy anther-specific gene with a male sterile Arabidopsis knockout phenotype is also in the Y-specific region, supporting a two-gene model for sex chromosome evolution.

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The asparagus genome sheds light on the origin and evolution of a young y chromosome

Dioecy, the presence of male and female flowers on separate individuals, is both widespread and uncommon within flowering plants, with only a few percent of dioecious species spread across most major phylogenetic taxa. It is therefore safe to assume that dioecy evolved independently in these different groups, which allows us to ask questions regarding the molecular and developmental mechanisms underlying these independent transitions to dioecy. We start this review by examining the problem from the standpoint of a genetic engineer trying to develop dioecy, discuss various potential solutions, and compare them to models proposed in the past and based on genetic and evolutionary considerations. Next, we present recent information regarding candidate sex determinants in three species, acquired using newly established genomic approaches. Although such specific information is still scarce, it is slowly becoming apparent that various genes or pathways can be altered to evolve dioecy.

One Hundred Ways to Invent the Sexes: Theoretical and Observed Paths to Dioecy in Plants.

Sex chromosomes have evolved independently in phylogenetically diverse flowering plant lineages. The genes governing sex determination in dioecious species remain unknown, but theory predicts that the linkage of genes influencing male and female function will spur the origin and early evolution of sex chromosomes. For example, in an XY system, the origin of an active Y may be spurred by the linkage of female suppressing and male promoting genes. Garden asparagus (Asparagus officinalis) serves as a model for plant sex chromosome evolution, given that it has recently evolved an XX/XY sex chromosome system. In order to elucidate the molecular basis of gender differences and sex determination, we used RNA-sequencing (RNA-Seq) to identify differentially expressed genes between female (XX), male (XY) and supermale (YY) individuals. We identified 570 differentially expressed genes, and showed that significantly more genes exhibited male-biased than female-biased expression in garden asparagus. In the context of anther development, we identified genes involved in pollen microspore and tapetum development that were specifically expressed in males and supermales. Comparative analysis of genes in the Arabidopsis thaliana, Zea mays and Oryza sativa anther development pathways shows that anther sterility in females probably occurs through interruption of tapetum development before microspore meiosis.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.13389

Sex-biased gene expression in dioecious garden asparagus (Asparagus officinalis).

Hundreds of land plant lineages have independently evolved separate sexes in either gametophytes (dioicy) or sporophytes (dioecy), but 43% of all dioecious angiosperms are found in just 34 entirely dioecious clades, suggesting that their mode of sex determination evolved a long time ago. Here, we review recent insights on the molecular mechanisms that underlie the evolutionary change from individuals that each produce male and female gametes to individuals specializing in the production of just one type of gamete. The canonical model of sex chromosome evolution in plants predicts that two sex-determining genes will become linked in a sex-determining region (SDR), followed by expanding recombination suppression, chromosome differentiation and, ultimately, degeneration. Experimental work, however, is showing that single genes function as master regulators in model systems, such as the liverwort Marchantia and the angiosperms Diospyros and Populus. In Populus, this type of regulatory function has been demonstrated by genome editing. In other systems, including Actinidia, Asparagus and Vitis, two coinherited factors appear to independently regulate female and male function, yet sex chromosome differentiation has remained low. We discuss the best-understood systems and evolutionary pathways to dioecy, and present a meta-analysis of the sizes and ages of SDRs. We propose that limited sexual conflict explains why most SDRs are small and sex chromosomes remain homomorphic. It appears that models of increasing recombination suppression with age do not apply because selection favours mechanisms in which sex determination depends on minimal differences, keeping it surgically precise.

Plant sex chromosomes defy evolutionary models of expanding recombination suppression and genetic degeneration

Garden asparagus (Asparagus officinalis L.) is an economically important plant with 2n = 2x = 20 chromosomes and a haploid genome size of 1323 Mb (Bennett and Leitch 2003). The karyotype of asparagus consists of five long (L), one medium (M) and four small (S) chromosomes (Löptien 1976). Plants of this dioecious species differ in producing either male or female flowers. The sexual dimorphism in asparagus is controlled by a region located on a pair of homomorphic sex chromosomes (chromosome L5) (Löptien 1979) termed the M locus (Flory 1932; Uno et al. 2002). Maleness is dominant with males normally heterogametic (Mm) and females homogametic (mm). Supermales (MM) can be produced from anther culture or from rarely occurring andromonoecious plants. The identification of chromosome L5 is important for studying sex chromosome evolution in asparagus. However, it was observed that the sex chromosome pair is homomorphic: X and Y chromosomes do not differ in morphology. Telgmann-Rauber et al. (2007) tried to clone the region determining sex in asparagus from its position in the genome. Molecular cytogenetic and sequence analyses of bacterial artificial chromosomes (BACs) flanking M locus indicate that the BAC contain highly repetitive sequences that localize to centromeric and pericentromeric locations on all asparagus chromosomes. However, the L5 chromosome could not be distinguished by BAC-FISH (Telgmann-Rauber et al. 2007). Recently, FISH has become a powerful and useful tool for the direct detection of specific DNA in genome. In this technique, ribosomal DNA genes (45S and 5S rDNA) are commonly used as markers for the physical mapping of plant chromosomes to analyse genomic organization (Nakayama et al. 2001; Lan et al. 2006; Grabowska-Joachimiak et al.

Karyotype of asparagus by physical mapping of 45S and 5S rDNA by FISH.

Male and female spinach (Spinacia oleracea L, 2n = 12) were karyotyped by chromosome measurements and subjected to fluorescence in situ hybridization with 45s rDNA The chromosome complement comprised two long nearly subtelocentrics, two short subtelocentrics, two acrocentrics, and six submetacentrics. The X and Y chromosomes were not distinguished by morphological analysis; thus, six obvious 45s rDNA sites were labeled with Texas red and clearly observed as red signals on three pairs of chromosomes. Among the three pairs of signal sites, a pair of the largest one was located on the short arms of chromosome 5, which is a satellite chromosome. Another prominent signal site was found on the short arms of chromosomes 2 and 6. However, from single images with 45s rDNA (red) of male or female spinach, weak 45s rDNA sites labeled with Texas red were observed on chromosomes 1 and 3. The weak 45s rDNA signals site were located on the large arm of chromosome 3 of both male and female spinach. Interestingly, differe...

https://www.tandfonline.com/doi/pdf/10.1080/00087114.2012.760879

Identification of sex chromosome of spinach by physical mapping of 45s rDNAs by FISH

Spinach has long been used as a model for genetic and physiological studies of sex determination and expression Although trisomic analysis from a cross between diploid and triploid plants identified the XY chromosome as the largest chromosome, no direct evidence has been provided to support this at the molecular level In this study, the largest chromosomes of spinach from mitotic metaphase spreads were microdissected using glass needles Degenerate oligonucleotide primed polymerase chain reaction was used to amplify the dissected chromosomes The amplified products from the Y chromosome were identified using the male-specific marker T11A For the first time, the largest spinach chromosome was confirmed to be a sex chromosome at the molecular level PCR products from the isolated chromosomes were used in an in situ probe mixture for painting the Y chromosome The fluorescence signals were mainly distributed on all chromosomes and four pair of weaker punctate fluorescence signal sites were observed on the terminal region of two pair of autosomes These findings provide a foundation for the study of sex chromosome evolution in spinach

Microdissection and painting of the Y chromosome in spinach (Spinacia oleracea)

The genome of spinach single chromosome complement is about 1000 Mbp, which is the model material to study the molecular mechanisms of plant sex differentiation. The cytological study showed that the biggest spinach chromosome (chromosome 1) was taken as spinach sex chromosome. It had three alleles of sex-related X,X(m) and Y. Many researchers have been trying to clone the sex-determining genes and investigated the molecular mechanism of spinach sex differentiation. However,there are no successful cloned reports about these genes. A new technology combining chromosome microdissection with hybridization-specific amplification (HSA) was adopted. The spinach Y chromosome degenerate oligonucleotide primed-PCR (DOP-PCR) products were hybridized with cDNA of the male spinach flowers in florescence. The female spinach genome was taken as blocker and cDNA library specifically expressed in Y chromosome was constructed. Moreover, expressed sequence tag (EST) sequences in cDNA library were cloned, sequenced and bioinformatics was analysed. There were 63 valid EST sequences obtained in this study. The fragment size was between 53 and 486 bp. BLASTn homologous alignment indicated that 12 EST sequences had homologous sequences of nucleic acids, the rest were new sequences. BLASTx homologous alignment indicated that 16 EST sequences had homologous protein-encoding nucleic acid sequence. The spinach Y chromosome-specific EST sequences laid the foundation for cloning the functional genes, specifically expressed in spinach Y chromosome. Meanwhile, the establishment of the technology system in the research provided a reference for rapid cloning of other biological sex chromosome-specific EST sequences.

Ying Cao

Papers

Karyotype of asparagus by physical mapping of 45S and 5S rDNA by FISH.

Identification of sex chromosome of spinach by physical mapping of 45s rDNAs by FISH

Microdissection and painting of the Y chromosome in spinach (Spinacia oleracea)

Rapid cloning and bioinformatic analysis of spinach Y chromosome-specific EST sequences.