Nature - USA (2020-01-02)

Nature | Vol 577 | 2 January 2020 | 83

related to immunity and stress responses in N. colorata, including
genes encoding nucleotide-binding leucine-rich repeat (NLR) proteins,
protein kinases and WRKY transcription factors, compared with those
in Amborella and some mesangiosperms (Extended Data Fig. 9, Sup-
plementary Note 8). It is possible that increased numbers of these genes
enabled water lilies to adapt to various ecological habitats globally.
In conclusion, the N. colorata genome offers a reference for compara-
tive genomics and for resolving the deep phylogenetic relationships
among the ANA-grade and mesangiosperms. It has also revealed a WGD
specific to Nymphaeales, and provides insights into the early evolution
of angiosperms on key innovations such as flower development and
floral scent and colour.

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availability are available at https://doi.org/10.1038/s41586-019-1852-5.

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a b

PetalStamenSepalJuvenile leafJuvenile leafstalkRootMature leafCarpelMature leafstalkJuvenile flower

NC11G0120830

NC7G0246550

NC2G0007900

NC11G0240820

NC11G0240810

NC13G0302010

NC12G0138630

NC11G0138610

NC11G0138620

NC2G0007880

NC11G0120810

NC11G0120820

200

400

600

800

1,000

1,200

1,400

d

e

FPKM

5.0

4.0

2.0

3.0

1.0

15 20 25

0

Abundance (total ion)

Retention time (min)

1

2

IS

3

456

78

9

10

11

11 fragrance molecules

1

3

56

8

c

Fatty acids

Relative activity

Butanoic

acid (C4)

Hexanoic

acid (C6)

Octanoic

acid (C8)

Decanoic

acid (C10)

Dodecanoic

acid (C12)

Tetradecanoic

acid (C14)

85 TPS-b

in N. colorata

Methyl

octanoate

m-Anisaldehyde

Methyl

decanoate

α-Bergamotene

Germacrene B

(E)-β-

Fanesene

Pentadecane

Isodaucene

Tetradecyne

Heptadecene

Farnesane

4

Monocots

Eudicots

Nymphaeales

Amborellales

Floral methyl decanoate

+

• 
  

Compound Gene

DAMT

Floral sesquiterpenes

+

+

+

• 
  

Compound Gene

TPS-a

TPS-a

TPS-b/g

• 
  


• 
  

Nonyl acetate

0

0.2

0.4

0.6

0.8

1.0

SABATH gene family

NC11G0120830

0.6

Marchantia polymorpha

Physcomitrella patens

Arabidopsis thaliana

Populus trichocarpa

Oryza sativa

Sorghum bicolor

Nymphaea colorata

Amborella trichopoda

Picea abies

Selaginella moellendorffii

Fig. 4 | Floral scent and biosynthesis in N. colorata. a, Gas chromatogram of
f loral volatiles from the f lower of N. colorata. The internal standard (IS) is nonyl
acetate. Methyl esters are in blue; terpenes are in red. Floral scent was
measured three times independently with similar results. b, Phylogenetic tree
of terpene synthases from N. colorata and representative plants showing the
subfamilies from a–h and x. c, Expression analysis of S A B AT H genes of
N. colorata showed that NC11G0120830 had the highest expression level in petal.

d, Relative activity of Escherichia coli-expressed NC11G0120830 with six fatty

acids as substrates, with the activity on decanoic acid set at 1.0. Data are

mean ± s.d. of three independent measurements. e, The presence (+) and

absence (−) of sesquiterpenes and methyl decanoate as f loral scent compounds

and their respective biosynthetic genes in four major lineages of angiosperms

when known. DAMT, decanoic acid methyltranferase.