1 Alternaria brassicae, a necrotrophic pathogen, causes Alternaria Leaf Spot, one of the 2 economically important diseases of Brassica crops. Many other Alternaria spp. such as A. brassicicola and 3A. alternata are known to cause secondary infections in the A. brassicae-infected Brassicas. The genome 4 architecture, pathogenicity factors, and determinants of host-specificity of A. brassicae are unknown. In 5 this study, we annotated and characterised the recently announced genome assembly of A. brassicae and 6 compared it with other Alternaria spp. to gain insights into its pathogenic lifestyle. Additionally, we 7 sequenced the genomes of two A. alternata isolates that were co-infecting B. juncea. Genome alignments 8 within the Alternaria spp. revealed high levels of synteny between most chromosomes with some 9 intrachromosomal rearrangements. We show for the first time that the genome of A. brassicae, a large-10 spored Alternaria species, contains a dispensable chromosome. We identified 460 A. brassicae-specific 11 genes, which included many secreted proteins and effectors. Furthermore, we have identified the gene 12 clusters responsible for the production of Destruxin-B, a known pathogenicity factor of A. brassicae. The 13 study provides a perspective into the unique and shared repertoire of genes within the Alternaria genus and 14 identifies genes that could be contributing to the pathogenic lifestyle of A. brassicae. 15 16 crops. Some of the most damaging species include Alternaria brassicae, A. brassicicola, A. 43 alternata, A. raphani, A. japonicus, and A. tenuissima. A. brassicae preferentially infects the oleiferous 44 Brassicas while the others are more devastating on the vegetable Brassicas. A. brassicae is particularly 45 more damaging in the hilly regions of the Indian subcontinent, where conducive climatic conditions allow 46 it to profusely reproduce and cause infections on almost all parts of the plant. Extensive screening for 47 resistance to A. brassicae in the cultivated Brassica germplasms has not revealed any source of resistance 2 .
48The factors that contribute to the pathogenicity of A. brassicae are relatively unknown.
49Pathogenicity of many Alternaria spp. has been mainly attributed to the secretion of host-specific toxins 50 (HSTs). HSTs induce pathogenesis on a rather narrow species range and are mostly indispensable for 51 pathogenicity. At least 12 A. alternata pathotypes have been reported to produce HSTs and thereby cause 52 disease on different species 3 . Many of the HST producing genes/gene clusters have been found on 53 supernumerary chromosomes or dispensable chromosomes 4 . A. brassicae has been reported to produce low 54 molecular weight cyclic depsipeptides named destruxins. Destruxin B is known to be a major phytotoxin 55 and is reported to be a probable HST of A. brassicae 5 . Additionally, a proteinaceous HST (ABR-toxin), 56 was isolated from the spore germination fluid of A. brassicae but was only partially characterised 6,7 .
57Genome sequencing and comparative analysi...