Abstract
Flowering, the transition from the vegetative to the generative phase, is a decisive time point in the lifecycle of a plant. Flowering is controlled by a complex network of transcription factors, photoreceptors, enzymes and miRNAs. In recent years, several studies gave rise to the hypothesis that this network is also strongly involved in the regulation of other important lifecycle processes ranging from germination and seed development through to fundamental developmental and yield-related traits. In the allopolyploid crop species Brassica napus, (genome AACC), homoeologous copies of flowering time regulatory genes are implicated in major phenological variation within the species, however the extent and control of intraspecific and intergenomic variation among flowering-time regulators is still unclear. To investigate differences among B. napus morphotypes in relation to flowering-time gene variation, we performed targeted deep sequencing of 29 regulatory flowering-time genes in four genetically and phenologically diverse B. napus accessions. The genotype panel included a winter-type oilseed rape, a winter fodder rape, a spring-type oilseed rape (all B. napus ssp. napus) and a swede (B. napus ssp. napobrassica), which show extreme differences in winter-hardiness, vernalization requirement and flowering behavior. A broad range of genetic variation was detected in the targeted genes for the different morphotypes, including non-synonymous SNPs, copy number variation and presence-absence variation. The results suggest that this broad variation in vernalization, clock and signaling genes could be a key driver of morphological differentiation for flowering-related traits in this recent allopolyploid crop species.
Highlights
As a recent allopolyploid species, Brassica napus L. is a very interesting model to investigate polyploidization and adaptation during crop evolution
In all other genes, including gene copies assigned to vernalization (Bna.EARLY FLOWERING IN SHORT DAYS (EFS), Bna.EARLY FLOWERING 7 (ELF7), Bna.FLOWERING LOCUS C (FLC), Bna.FRI, Bna.SUPPRESSOR OF FRIGIDA 4 (SUF4), Bna.TERMINAL FLOWER 2 (TFL2), Bna.VERNALIZATION INSENSITIVE 3 (VIN3)), photoperiod (Bna.CO, Bna.CO-like, Bna.CRYPTOCHROME 2 (CRY2) and Bna.PHYTOCHROME A (PHYA)), gibberellin (Bna.GA3ox1), clock (Bna.CYCLING DOF FACTOR 1 (CDF1), Bna.EARLY FLOWERING 3 (ELF3), Bna.ZTL) and signaling (Bna.AGAMOUS-LIKE 24 (AGL24), Bna.APETALA 1 (AP1), Bna.FLOWERING LOCUS D (FD), Bna.FLOWERING LOCUS T (FT), Bna.FUL, Bna.LFY, Bna.SQUAMOSA PROMOTER BINDING PROTEIN-LIKE 3 (SPL3), Bna.SUPPRESSOR OF CONSTANS 1 (SOC1), Bna.TEMPRANILLO 1 (TEM1), and Bna.TERMINAL FLOWER 1 (TFL1)), we found potentially functional amino acid modifications in the gene products of at least one homolog/paralog within the four different B. napus morphotypes (Figure 4)
We found further mutations in gene copies from the photoperiod pathway (Bna.CO, Bna.CO-like 2, Bna.PHYA, Bna.PHYTOCHROME B (PHYB)), gibberellin synthesis (Bna.GA3ox1), temperature signaling (Bna.SPL3, Bna.ELF3), the central signaling molecules (Bna.FT, Bna.FD) and downstream effectors (Bna.AGL24, Bna.AP1, Bna.FUL, Bna.SOC1, Bna.TFL1)
Summary
No wild forms of B. napus are known, and intensive selection and breeding following its anthropogenically-influenced polyploidization has led to cultivation of very different phenological types. This has caused the diversification of distinct gene pools adapted to highly different eco-geographic zones of Europe, Asia/Australia and North America. Very early-flowering morphotypes, without vernalization requirement, are today widely grown in Canada (as canola) and northern Europe (as spring oilseed rape), where harsh winters prohibit autumn-sown crops. Later-flowering “semi-winter” oilseed forms, requiring only mild vernalization, are prevalent in China and Australia, while autumn-sown oilseed rape is today the most important oilseed crop in temperate regions of Europe (Friedt and Snowdon, 2010). Swedes generally have a strong vernalization requirement but tend to lack the strong winter-hardiness of winter oilseed rape (Friedt and Snowdon, 2010)
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