The most complete published dataset for this question comes from a genome-wide survey of 1,087 cyclin genes across ten Brassicaceae genomes (six Brassica species of U's triangle plus four Arabidopsis species), which used 49 curated Arabidopsis thaliana cyclins as queries, Pfam domain validation, and reciprocal BLASTp, then OrthoFinder to assign one-to-one orthologs between each diploid progenitor and its allotetraploid derivative, with retention defined as an ortholog located on the expected subgenome (e.g., B. rapa genes on the A subgenome) and "exchange" defined as an ortholog found on a non-expected subgenome .
The diploid progenitor reference counts and allotetraploid totals needed for a paired bar chart ("retention capacity" β what the allotetraploid starts with before fractionation) are published:
| Species | Ploidy / genome | Total cyclin genes (23 types, BLASTp+Pfam, 49 At queries) | Duplication events (genes involved) |
|---|---|---|---|
| B. rapa | AA (diploid) | 76β90* | 154 (87) |
| B. oleracea | CC (diploid) | ~90 | 140 (94) |
| B. nigra | BB (diploid) | ~90 | <160 |
| B. napus | AACC (allotetraploid) | 189 | 372 (186) |
| B. juncea | AABB (allotetraploid) | 185 | 377 (185) β highest |
| B. carinata | BBCC (allotetraploid) | ~166 | 299 (167) |
*76 is the independent B. rapa genome-wide count using BrassicaDB annotation with 10 Arabidopsis cyclin queries; ~90 is from the 10-species comparative framework (diploid average 71.5, range 90β189 across all six Brassica) β the discrepancy reflects differing annotation sets and query panels .
Within B. napus, a CYCB-specific study found 25 CYCB genes in the Darmor-bzh reference (11 on the A subgenome, 13 on the C subgenome, 1 on a scaffold), expanding to 299 CYCB genes across 11 accessions (20β30 typical, up to 50 in Ningyou) β showing accession-level copy-number variation on top of the fixed subgenome-level retention . The near-even 11:13 A:C CYCB split is consistent with the balanced (~2:2) expectation from subgenome-wide fractionation analyses where multi-copy genes show ~40% per subgenome and no strong subgenome bias .
| Allotetraploid | Progenitor (genome) | Expected subgenome | Ortholog pairs | Retained (n) | Retention % | Exchanged % |
|---|---|---|---|---|---|---|
| B. napus (AACC) | B. rapa (A) | A | 127 | 86 | 67.7 | 32.2 (βC) |
| B. napus (AACC) | B. oleracea (C) | C | 125 | 86 | 68.8 | 31.2 (βA) |
| B. juncea (AABB) | B. rapa (A) | A | 115 | 85 | 73.9 | 26.0 (βB) |
| B. juncea (AABB) | B. nigra (B) | B | 96 | 84 | 87.5 | 12.5 (βA) |
| B. carinata (BBCC) | B. nigra (B) | B | 81 | 54 | 66.6 | 33.3 (βC) |
| B. carinata (BBCC) | B. oleracea (C) | C | 98 | 69 | 70.4 | 29.5 (βB) |
All values from the published ortholog-retention analysis (Table S9 summarized in main text); retention ranged 66.6β87.5% and exchange 12.5β33.3% across the six progenitorβallotetraploid combinations .
A key asymmetry: the B genome in B. juncea shows the highest retention (87.5%), while the same B. nigra progenitor shows the lowest retention in B. carinata (66.6%) β retention is not an intrinsic property of the progenitor genome but depends on the recipient allotetraploid background, consistent with extensive homoeologous exchange and genome reshuffling in allopolyploid Brassica .
X-axis: progenitorβallotetraploid combination (6 categories). Y-axis: % retained on expected subgenome. Overlay line: % exchanged.
combinations = {
"B.rapaβB.napus (A)": 67.7, "B.oleraceaβB.napus (C)": 68.8,
"B.rapaβB.juncea (A)": 73.9, "B.nigraβB.juncea (B)": 87.5,
"B.nigraβB.carinata (B)": 66.6, "B.oleraceaβB.carinata (C)": 70.4}
exchanged = {"A":32.2,"C":31.2,"A":26.0,"B":12.5,"B":33.3,"C":29.5}
Paired bars per U's-triangle progeny: progenitor counts vs allotetraploid counts (e.g., B. rapa 76β90 / B. oleracea ~90 vs B. napus 189; B. rapa / B. nigra vs B. juncea 185; B. nigra / B. oleracea vs B. carinata ~166), with a reference line at the diploid average 71.5.
Per-cyclin-type ortholog data for a heatmap (ortholog pairs, progenitor genes involved): cycT1 (83 pairs, 38 genes), cycA2 (81, 36), cycB2 (76, 34), cycB1 (69, 32), cycD3 (64, 28), cycU4 (61, 24), cycA3 (55, 24), cycA1 (48, 19); among D-types, cycD3 retained the most pairs (64) while cycD5 was lowest (14) . Duplication-mode breakdown for a proportional bar chart: of 1,845 total duplication events, 969 were WGD, 730 dispersed, 75 transposed, 51 tandem, 20 proximal; intra- vs inter-genomic splits in allotetraploids were B. napus 149 intra (83 A + 66 C) vs 214 inter; B. juncea 147 intra vs 212 inter; B. carinata 120 intra vs 154 inter .
1. Retention is high but incomplete and background-dependent (Fig. A). Cyclin ortholog retention on expected subgenomes spans 66.6β87.5% across the three Brassica allotetraploids; homoeologous exchange (12.5β33.3%) is universal.
2. Allotetraploids exceed the simple additive expectation (Fig. B). B. napus (189) and B. juncea (185) hold substantially more cyclins than the sum of typical diploid progenitors would minimally predict (~2Γ71.5 = 143), reflecting post-polyploidization duplications (especially WGD-type events) alongside retention; B. carinata (~166) retains the fewest.
3. Mechanistic drivers: the shared Brassica whole-genome triplication and subsequent allopolyploidization supplied the raw copy number , while differential homoeolog gene expression and dosage-balance selection then shaped which copies were retained .
For flowering-time relevance in B. napus: two cyclins (Bna21cycA2, Bna113cycD4) co-localize with flowering-time QTL and carry diagnostic amino-acid polymorphisms in early-flowering cultivars (Westar, No2127) β a candidate downstream application of the retention data .
Explore B. napus cyclin subgenome data β Explore B. juncea / B. carinata retention β Explore B. rapa cyclin family β
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