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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Any methods, additional references, Nature Research reporting sum-
maries, source data, extended data, supplementary information,
acknowledgements, peer review information; details of author con-
tributions and competing interests; and statements of data and code
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.