Cell - 8 September 2016

confirmed by the average per-site nucleotide divergence (dxy),
which is significantly lower between Britain and the United States
(averagedxy= 1.97 310 ^3 ) than between Belgium/Germany and
the United States strains (averagedxy= 2.26 310 ^3 ) (Wilcoxon

signed-rank test, p < 0.001) (Table S8). This suggests that the

origin of the United States brewing strains can be traced back

to the introduction of beer culture in the United States by early

17 thcentury British settlers (Van Wieren, 1995). Third, in contrast

B

C

A

Britain US Bel/Ger Beer 2 Mixed Wine Asia Mosaic

0

25

50

75

100

Sporulation efficiency (%)

0

25

50

75

100

Spore viability (%)

Britain US Bel/Ger Beer 2 Mixed Wine Asia Mosaic

I II III IV V VI VII VIII IX X XI XII XIII XIV XV XVI

0.50

1.00 Britain

US

Bel/Ger

Mixed

Wine

Asia

0.00

0.50

1.00

0.00

0.50

1.00

0.00

0.50

1.00

0.00

0.50

1.00

0.00

0.50

1.00

0.00

0.50

1.00

0.00

Level of heterozygosity

Spore viability (%) Sporulation efficiency (%)

Nr. of heterozygous SNPs (x10

3 )

Nr of heterozygous SNPs (x10

3 ) R^2 = 0.17, p < 0.001 R^2 = 0.03, p = 0.022

0

20

40

60

80

0 25 50 75 100

0

20

40

60

80

0 25 50 75 100

0

20

40

60

0 25 50 75 100

R^2 = 0.19, p < 0.001

Sporulation efficiency (%)

0

20

40

60

0 25 50 75 100

R^2 = 0.16, p < 0.001

Spore viability (%)

Amplified/Deleted genome fraction (%)

D E

F G

Beer 2

Britain

US

Bel/Ger

Beer 2

Mixed

Wine

Asia

Mosaic

Britain

US

Bel/Ger

Beer 2

Mixed

Wine

Asia

Mosaic

Britain

US

Bel/Ger

Beer 2

Mixed

Wine

Asia

Mosaic

Britain

US

Bel/Ger

Beer 2

Mixed

Wine

Asia

Mosaic

Amplified/Deleted genome fraction (%)

Figure 4. The Reproductive Lifestyle of IndustrialS. cerevisiaeStrains
(A) Violin plots depicting sporulation efficiency of all strains from different subpopulations.
(B) Violin plots depicting spore viability of all sporulating strains from different subpopulations.
(C) Visualization of the level of heterozygosity across the genome of the different subpopulations, calculated as the ratio of heterozygous/homozygous SNPs in
10 kb windows.
(D–G) Scatter plots depicting the correlation between the number of heterozygous loci and spore viability (D) or sporulation efficiency (E), and the correlation
between the fraction of the genome subjected to large (>20 kb) structural variation and spore viability (F) or sporulation efficiency (G). Dot colors indicate
subpopulations similar to the color code ofFigure 1B.
See alsoFigure S4.

Cell 166 , 1397–1410, September 8, 2016 1405