Nelumbo: Difference between revisions

Nelumbo; Temporal range: Albian to present PreꞒ Ꞓ O S D C P T J K Pg N
	Nelumbo nucifera (sacred lotus)
	Nelumbo lutea (American lotus)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Order:	Proteales
Family:	Nelumbonaceae
Genus:	Nelumbo; Adans.
Species
	Nelumbo lutea; Nelumbo nucifera; †Nelumbo aureavallis; †Nelumbo changchangensis; †Nelumbo minima; †Nelumbo nipponica; †Nelumbo orientalis; †Nelumbo protolutea; 30+ more from fossil remains;

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Nelumbo /nɪˈlʌmboʊ/^[2] is a genus of aquatic plants with large, showy flowers. Members are commonly called lotus, though the name is also applied to various other plants and plant groups, including the unrelated genus Lotus. Members outwardly resemble those in the family Nymphaeaceae ("water lilies"), but Nelumbo is actually very distant from that family.

Nelumbo is an ancient genus, with dozens of species known from fossil remains since the Early Cretaceous. However, there are only two known living species of lotus. One is the better-known Nelumbo nucifera, which is native to East Asia, South Asia, Southeast Asia, and probably Australia and is commonly cultivated for consumption and use in traditional Chinese medicine. The other lotus is Nelumbo lutea, which is native to North America and the Caribbean. Horticultural hybrids have been produced between these two allopatric species.

Description

Ultrahydrophobicity

The leaves of Nelumbo are highly water-repellent (i.e. they exhibit ultrahydrophobicity) and have given the name to what is called the lotus effect.^[3] Ultrahydrophobicity involves two criteria: a very high water contact angle between the droplet of water and the leaf surface, and a very low roll-off angle.^[4] This means that the water must contact the leaf surface at exactly one, minuscule point, and any manipulation of the leaf by changing its angle will result in the water droplet rolling off of the leaf.^[4] Ultrahydrophobicity is conferred by the usually dense layer of papillae on the surface of the Nelumbo leaves, and the small, robust, waxy tubules that protrude off each papilla.^[5] This helps reduce the area of contact between the water droplet and the leaf.^[5]

Ultrahydrophobicity is said to confer a very important evolutionary advantage. As an aquatic plant with leaves that rest on the water's surface, the genus Nelumbo is characterized by its concentration of stomata on the upper epidermis of its leaves, unlike most other plants which concentrate their stomata on the lower epidermis, underneath the leaf.^[5] The collection of water on the upper epidermis, whether that be by rain, mist, or the nearby disturbance of water, is very detrimental to the leaf's ability to perform gas exchange through its stomata. Thus, Nelumbo's ultrahydrophobicity allows the water droplets to accumulate together very quickly, and then roll off of the leaf very easily at the slightest disturbance of the leaf, a process which allows its stomata to function normally without restriction due to blockage by water droplets.^[6]

Thermoregulation

An uncommon property of the genus Nelumbo is that it can generate heat,^[7] which it does by using the alternative oxidase pathway (AOX).^[8]^[9] This pathway involves a different, alternative exchange of electrons from the usual pathway that electrons follow when generating energy in mitochondria, known as the AOX, or alternative oxidase pathway.

The typical pathway in plant mitochondria involves cytochrome complexes. The pathway used to generate heat in Nelumbo involves cyanide-resistant alternative oxidase, which is a different electron acceptor than the usual cytochrome complexes.^[10] The plant also reduces ubiquitin concentrations while in thermogenesis, which allows the AOX in the plant to function without degradation.^[11] Thermogenesis is restricted to the receptacle, stamen, and petals of the flower, but each of these parts produce heat independently without relying on the heat production in other parts of the flower.^[12]

There are several theories about the function of thermogenesis, especially in an aquatic genus such as Nelumbo. The most common theory posits that thermogenesis in flowers attracts pollinators, for a variety of reasons. Heated flowers may attract insect pollinators. As the pollinators warm themselves while resting inside the flower, they deposit and pick up pollen onto and from the flower.^[7] The thermogenic environment might also be conducive to pollinator mating; pollinators may require a certain temperature for reproduction. By providing an ideal thermogenic environment, the flower is pollinated by mating pollinators.^[13] Others theorize that heat production facilitates the release of volatile compounds into the air to attract pollinators flying over water, or that the heat is recognizable in the dark by thermo-sensitive pollinators. None have been conclusively proven to be more plausible than the others.^[14]

After anthesis, the receptacle of the lotus transitions from a primarily thermogenic to a photosynthetic structure, as seen in the rapid and dramatic increase in photosystems, photosynthetically involved pigments, electron transport rates, and the presence of 13^C in the receptacle and petals, all of which assist in increasing photosynthesis rates.^[15] After this transition, all thermogenesis in the flower is lost. Pollinators do not need to be attracted once the ovary is fertilized, and thus the receptacle's resources are better used when it is photosynthesizing to produce carbohydrates that can increase plant biomass or fruit mass.^[13]

Other plants utilize thermoregulation in their life cycles. Among these is the eastern skunk cabbage, which heats itself to melt any ice above it, and push through the ground in early spring.^[16] Also, the elephant yam, which heats its flowers to attract pollinators. In addition, the carrion flower, which heats itself to disperse water vapor through the air, carrying its scent further, thus attracting more pollinators.

Similar species

The leaves of Nelumbo can be distinguished from those of genera in the family Nymphaeaceae as they are peltate, that is they have fully circular leaves. Nymphaea, on the other hand, has a single characteristic notch from the edge in to the center of the lily pad. The seedpod of Nelumbo is very distinctive.

Taxonomy

Taxonomic history

The Cronquist system of 1981 recognizes the family Nelumbonaceae but places it in the water lily order Nymphaeales. The Dahlgren system of 1985 and Thorne system of 1992 both recognize the family and place it in its own order, Nelumbonales. The United States Department of Agriculture still classifies the lotus family within the water lily order.^[17]

There is residual disagreement over which family the genus should be placed in. Traditional classification systems recognized Nelumbo as part of the Nymphaeaceae, but traditional taxonomists were likely misled by convergent evolution associated with an evolutionary shift from a terrestrial to an aquatic lifestyle. In the older classification systems it was recognized under the order Nymphaeales or Nelumbonales.

Modern classification

Nelumbo is currently recognized as the only living genus in Nelumbonaceae, one of several distinctive families in the eudicot order of the Proteales. Its closest living relatives, the (Proteaceae and Platanaceae), are shrubs or trees.

The APG IV system of 2016 recognizes Nelumbonaceae as a distinct family and places it in the order Proteales in the eudicot clade, as do the earlier APG III and APG II systems.^[18]

Phylogeny

There are several fossil species known from Cretaceous, Paleogene and Neogene aged strata throughout Eurasia and North America. Despite the ancient origins of this genus and the wide geographic separation of the two extant species (N. nucifera and N. lutea), phylogenetic evidence indicates that they diverged rather recently, during the early Pleistocene (about 2 million years ago).^[19]

Species

Extant species

Nelumbo lutea Willd. – American lotus (Eastern United States, Mexico, Greater Antilles, Honduras)
Nelumbo nucifera Gaertn. – sacred or Indian lotus, also known as the Rose of India and the sacred water lily of Hinduism and Buddhism.^[20] It is the national flower of India and Vietnam. Its roots and seeds are also used widely in cooking in East Asia, South Asia and Southeast Asia.

Fossil species

Nearly 30 fossil species are known from the mid-Cretaceous to the present.^[1]

†Nelumbo aureavallis Hickey – Eocene (North Dakota), described from leaves found in the Golden Valley Formation in North Dakota, USA.^[21]
†Nelumbo changchangensis Eocene, (Hainan Island, China), described from several fossils of leaves, seedpods, and rhizomes from the Eocene-aged strata in the Changchang Basin, of Hainan Island.
†Nelumbo choffati Early Cretaceous (Portugal), leaves known from the Albian. One of the earliest known species.^[1]
†Nelumbo jiayinensis Late Cretaceous, (Heilongjiang, China), leaves described from the Santonian-aged Yong'ancun Formation^[22]
†Nelumbo lusitanica Early Cretaceous (Portugal), leaves known from the Albian. One of the earliest known species.^[1]
†Nelumbo minima Pliocene (Netherlands), described from leaves and seedpods that suggest a very small plant. Originally described as a member of the genus Nelumbites, as "Nelumbites minimus."
†Nelumbo nipponica Eocene-Miocene, fossil leaves are known from Eocene-aged strata in Japan, and Miocene-aged strata in Russia.
†Nelumbo orientalis Cretaceous (Japan), fossils found in Cretaceous-aged strata of Japan. The Sarao Formation, which they are known from, was formerly considered of Early Cretaceous age, but more recent studies support a Maastrichtian age for it.^[23]^[24]
†Nelumbo protolutea Eocene (Mississippi), fossils of leaves strongly suggest a plant similar in form to the American lotus.
†Nelumbo weymouthi Early Cretaceous (Wyoming, US), leaves known from the Albian. One of the earliest known species.^[1]

Etymology

The genus name is derived from Sinhala: නෙළුම් neḷum, the name for Nelumbo nucifera.^[25]

Uses

Vinegared lotus root slices with ginger and citron as eaten in Japan

The entire plant can be eaten either raw or cooked. The underwater portion is high in starch. The fleshy parts can be dug from the mud and baked or boiled. The young leaves can be boiled. The seeds are palatable and can be eaten raw or dried and ground into flour.^[26] The stem fibers are also used to make lotus silk.^[27]

Culture

The sacred lotus, N. nucifera, is sacred in both Hinduism and Buddhism.^[20] It is the floral emblem of both India and Vietnam.

References

^ ^a ^b ^c ^d ^e Li, Ya; Svetlana, Popova; Yao, Jianxin; Li, Chengsen (2014). "A Review on the Taxonomic, Evolutionary and Phytogeographic Studies of the Lotus Plant (Nelumbonaceae: Nelumbo )". Acta Geologica Sinica - English Edition. 88 (4): 1252–1261. Bibcode:2014AcGlS..88.1252L. doi:10.1111/1755-6724.12287. ISSN 1000-9515.
^ "nelumbo". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)
^ Darmanin T, Guittard F (1 June 2015). "Superhydrophobic and superoleophobic properties in nature". Materials Today. 18 (5): 273–285. doi:10.1016/j.mattod.2015.01.001.
^ ^a ^b Marmur A (2004-04-01). "The Lotus Effect: Superhydrophobicity and Metastability". Langmuir. 20 (9): 3517–3519. doi:10.1021/la036369u. PMID 15875376.
^ ^a ^b ^c Zhang Y, Wu H, Yu X, Chen F, Wu J (March 2012). "Microscopic Observations of the Lotus Leaf for Explaining the Outstanding Mechanical Properties". Journal of Bionic Engineering. 9 (1): 84–90. doi:10.1016/S1672-6529(11)60100-5. S2CID 137076244.
^ Ensikat HJ, Ditsche-Kuru P, Neinhuis C, Barthlott W (2011-03-10). "Superhydrophobicity in perfection: the outstanding properties of the lotus leaf". Beilstein Journal of Nanotechnology. 2: 152–61. doi:10.3762/bjnano.2.19. PMC 3148040. PMID 21977427.
^ ^a ^b Watling JR, Robinson SA, Seymour RS (April 2006). "Contribution of the alternative pathway to respiration during thermogenesis in flowers of the sacred lotus". Plant Physiology. 140 (4): 1367–73. doi:10.1104/pp.105.075523. PMC 1435819. PMID 16461386.
^ Seymour RS, Schultze-Motel P, Lamprecht I (1 July 1998). "Heat production by sacred lotus flowers depends on ambient temperature, not light cycle". Journal of Experimental Botany. 49 (324): 1213–1217. doi:10.1093/jxb/49.324.1213.
^ Grant NM, Miller RA, Watling JR, Robinson SA (2010-11-12). "Distribution of thermogenic activity in floral tissues of Nelumbo nucifera". Functional Plant Biology. 37 (11): 1085–1095. doi:10.1071/FP10024. ISSN 1445-4416.
^ Hiroma T, Ito K, Hara M, Torisu R (2011-06-01). "Analysis of the Lotus Thermoregulation System from the Perspective of Control Engineering". Shokubutsu Kankyo Kogaku (in Japanese). 23 (2): 52–58. doi:10.2525/shita.23.52. ISSN 1880-2028.
^ Wang R, Zhang Z (March 2015). "Floral thermogenesis: An adaptive strategy of pollination biology in Magnoliaceae". Communicative & Integrative Biology. 8 (1): e992746. doi:10.4161/19420889.2014.992746. PMC 4594551. PMID 26844867.
^ Li JK, Huang SQ (May 2009). "Flower thermoregulation facilitates fertilization in Asian sacred lotus". Annals of Botany. 103 (7): 1159–63. doi:10.1093/aob/mcp051. PMC 2707905. PMID 19282320.
^ ^a ^b Miller RE, Watling JR, Robinson SA (2009). "Functional transition in the floral receptacle of the sacred lotus (Nelumbo nucifera): from thermogenesis to photosynthesis". Functional Plant Biology. 36 (5): 471–480. doi:10.1071/FP08326. PMID 32688661. S2CID 54588650.
^ Wagner AM, Krab K, Wagner MJ, Moore AL (2008-07-01). "Regulation of thermogenesis in flowering Araceae: the role of the alternative oxidase". Biochimica et Biophysica Acta (BBA) - Bioenergetics. 1777 (7–8): 993–1000. doi:10.1016/j.bbabio.2008.04.001. PMID 18440298.
^ Grant NM, Miller RE, Watling JR, Robinson SA (2008-02-01). "Synchronicity of thermogenic activity, alternative pathway respiratory flux, AOX protein content, and carbohydrates in receptacle tissues of sacred lotus during floral development". Journal of Experimental Botany. 59 (3): 705–14. doi:10.1093/jxb/erm333. hdl:2440/52460. PMID 18252702.
^ Ito K, Ito T, Onda Y, Uemura M (2004-03-15). "Temperature-Triggered Periodical Thermogenic Oscillations in Skunk Cabbage (Symplocarpus foetidus)". Plant and Cell Physiology. 45 (3): 257–264. doi:10.1093/pcp/pch038. ISSN 0032-0781. PMID 15047873.
^ NRCS. "Nelumbo nucifera". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 2024-11-03.
^ Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. ISSN 0024-4074.
^ Wu, Zhihua; Gui, Songtao; Quan, Zhiwu; Pan, Lei; Wang, Shuzhen; Ke, Weidong; Liang, Dequan; Ding, Yi (2014-11-19). "A precise chloroplast genome of Nelumbo nucifera (Nelumbonaceae) evaluated with Sanger, Illumina MiSeq, and PacBio RS II sequencing platforms: insight into the plastid evolution of basal eudicots". BMC Plant Biology. 14 (1): 289. doi:10.1186/s12870-014-0289-0. ISSN 1471-2229. PMC 4245832. PMID 25407166.
^ ^a ^b "Nelumbo nucifera (sacred lotus)". Kew Royal Botanic Gardens. Archived from the original on 30 May 2014. Retrieved 26 July 2015.
^ Hickey L (1977). Stratigraphy and Paleobotany of the Golden Valley Formation (Early Tertiary) of Western North Dakota. Boulder, Colorado: Geological Society of America. pp. 110 & Plate 5. ISBN 978-0-8137-1150-8.
^ Liang, Fei; Sun, Ge; Yang, Tao; Bai, Shuchong (April 2018). "Nelumbo jiayinensis sp. nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China". Cretaceous Research. 84: 134–140. Bibcode:2018CrRes..84..134L. doi:10.1016/j.cretres.2017.11.007. ISSN 0195-6671.
^ "Discovery of Nelumbo from the Asuwa Flora (Upper Cretaceous) in Fukui Prefecture in the Inner Side of Central Japan". gbank.gsj.jp. Retrieved 2024-07-19.
^ 佐沢, 豪; ルグラン, ジュリアン; 矢部, 淳; 上松, 佐知子; 指田, 勝男 (2020). "福井県東部，池田町に分布する上部白亜系足羽層の年代-花粉層位学的アプローチ". 地質学雑誌. 126 (4): 215–221. doi:10.5575/geosoc.2020.0004.
^ Hyam R, Pankhurst RJ (1995). Plants and their names: a concise dictionary. Oxford: Oxford University Press. ISBN 978-0-19-866189-4.
^ The Complete Guide to Edible Wild Plants. United States Department of the Army. New York: Skyhorse Publishing. 2009. p. 64. ISBN 978-1-60239-692-0. OCLC 277203364.{{cite book}}: CS1 maint: others (link)
^ "Weaving success: Manipur woman seeks to employ more locals in her lotus silk venture". The Print. PTI. 11 September 2022.

External links

links at CSDL Archived 2008-10-12 at the Wayback Machine

[Li-2014-1] Li, Ya; Svetlana, Popova; Yao, Jianxin; Li, Chengsen (2014). "A Review on the Taxonomic, Evolutionary and Phytogeographic Studies of the Lotus Plant (Nelumbonaceae: Nelumbo )". Acta Geologica Sinica - English Edition. 88 (4): 1252–1261. Bibcode:2014AcGlS..88.1252L. doi:10.1111/1755-6724.12287. ISSN 1000-9515.

[2] "nelumbo". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)

[3] Darmanin T, Guittard F (1 June 2015). "Superhydrophobic and superoleophobic properties in nature". Materials Today. 18 (5): 273–285. doi:10.1016/j.mattod.2015.01.001.

[Marmur_2004-4] Marmur A (2004-04-01). "The Lotus Effect: Superhydrophobicity and Metastability". Langmuir. 20 (9): 3517–3519. doi:10.1021/la036369u. PMID 15875376.

[Zhang_2012-5] Zhang Y, Wu H, Yu X, Chen F, Wu J (March 2012). "Microscopic Observations of the Lotus Leaf for Explaining the Outstanding Mechanical Properties". Journal of Bionic Engineering. 9 (1): 84–90. doi:10.1016/S1672-6529(11)60100-5. S2CID 137076244.

[6] Ensikat HJ, Ditsche-Kuru P, Neinhuis C, Barthlott W (2011-03-10). "Superhydrophobicity in perfection: the outstanding properties of the lotus leaf". Beilstein Journal of Nanotechnology. 2: 152–61. doi:10.3762/bjnano.2.19. PMC 3148040. PMID 21977427.

[Watling_2006-7] Watling JR, Robinson SA, Seymour RS (April 2006). "Contribution of the alternative pathway to respiration during thermogenesis in flowers of the sacred lotus". Plant Physiology. 140 (4): 1367–73. doi:10.1104/pp.105.075523. PMC 1435819. PMID 16461386.

[8] Seymour RS, Schultze-Motel P, Lamprecht I (1 July 1998). "Heat production by sacred lotus flowers depends on ambient temperature, not light cycle". Journal of Experimental Botany. 49 (324): 1213–1217. doi:10.1093/jxb/49.324.1213.

[Grant_2010-9] Grant NM, Miller RA, Watling JR, Robinson SA (2010-11-12). "Distribution of thermogenic activity in floral tissues of Nelumbo nucifera". Functional Plant Biology. 37 (11): 1085–1095. doi:10.1071/FP10024. ISSN 1445-4416.

[Hiroma_2011-10] Hiroma T, Ito K, Hara M, Torisu R (2011-06-01). "Analysis of the Lotus Thermoregulation System from the Perspective of Control Engineering". Shokubutsu Kankyo Kogaku (in Japanese). 23 (2): 52–58. doi:10.2525/shita.23.52. ISSN 1880-2028.

[Wang_2015-11] Wang R, Zhang Z (March 2015). "Floral thermogenesis: An adaptive strategy of pollination biology in Magnoliaceae". Communicative & Integrative Biology. 8 (1): e992746. doi:10.4161/19420889.2014.992746. PMC 4594551. PMID 26844867.

[Li_2009-12] Li JK, Huang SQ (May 2009). "Flower thermoregulation facilitates fertilization in Asian sacred lotus". Annals of Botany. 103 (7): 1159–63. doi:10.1093/aob/mcp051. PMC 2707905. PMID 19282320.

[Miller_2009-13] Miller RE, Watling JR, Robinson SA (2009). "Functional transition in the floral receptacle of the sacred lotus (Nelumbo nucifera): from thermogenesis to photosynthesis". Functional Plant Biology. 36 (5): 471–480. doi:10.1071/FP08326. PMID 32688661. S2CID 54588650.

[Wagner_2008-14] Wagner AM, Krab K, Wagner MJ, Moore AL (2008-07-01). "Regulation of thermogenesis in flowering Araceae: the role of the alternative oxidase". Biochimica et Biophysica Acta (BBA) - Bioenergetics. 1777 (7–8): 993–1000. doi:10.1016/j.bbabio.2008.04.001. PMID 18440298.

[Grant_2008-15] Grant NM, Miller RE, Watling JR, Robinson SA (2008-02-01). "Synchronicity of thermogenic activity, alternative pathway respiratory flux, AOX protein content, and carbohydrates in receptacle tissues of sacred lotus during floral development". Journal of Experimental Botany. 59 (3): 705–14. doi:10.1093/jxb/erm333. hdl:2440/52460. PMID 18252702.

[16] Ito K, Ito T, Onda Y, Uemura M (2004-03-15). "Temperature-Triggered Periodical Thermogenic Oscillations in Skunk Cabbage (Symplocarpus foetidus)". Plant and Cell Physiology. 45 (3): 257–264. doi:10.1093/pcp/pch038. ISSN 0032-0781. PMID 15047873.

[17] NRCS. "Nelumbo nucifera". PLANTS Database. United States Department of Agriculture (USDA). Retrieved 2024-11-03.

[APGIV-18] Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. ISSN 0024-4074.

[19] Wu, Zhihua; Gui, Songtao; Quan, Zhiwu; Pan, Lei; Wang, Shuzhen; Ke, Weidong; Liang, Dequan; Ding, Yi (2014-11-19). "A precise chloroplast genome of Nelumbo nucifera (Nelumbonaceae) evaluated with Sanger, Illumina MiSeq, and PacBio RS II sequencing platforms: insight into the plastid evolution of basal eudicots". BMC Plant Biology. 14 (1): 289. doi:10.1186/s12870-014-0289-0. ISSN 1471-2229. PMC 4245832. PMID 25407166.

[Kew-20] "Nelumbo nucifera (sacred lotus)". Kew Royal Botanic Gardens. Archived from the original on 30 May 2014. Retrieved 26 July 2015.

[Hickey1977-21] Hickey L (1977). Stratigraphy and Paleobotany of the Golden Valley Formation (Early Tertiary) of Western North Dakota. Boulder, Colorado: Geological Society of America. pp. 110 & Plate 5. ISBN 978-0-8137-1150-8.

[22] Liang, Fei; Sun, Ge; Yang, Tao; Bai, Shuchong (April 2018). "Nelumbo jiayinensis sp. nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China". Cretaceous Research. 84: 134–140. Bibcode:2018CrRes..84..134L. doi:10.1016/j.cretres.2017.11.007. ISSN 0195-6671.

[23] "Discovery of Nelumbo from the Asuwa Flora (Upper Cretaceous) in Fukui Prefecture in the Inner Side of Central Japan". gbank.gsj.jp. Retrieved 2024-07-19.

[24] 佐沢, 豪; ルグラン, ジュリアン; 矢部, 淳; 上松, 佐知子; 指田, 勝男 (2020). "福井県東部，池田町に分布する上部白亜系足羽層の年代-花粉層位学的アプローチ". 地質学雑誌. 126 (4): 215–221. doi:10.5575/geosoc.2020.0004.

[HyamPank95-25] Hyam R, Pankhurst RJ (1995). Plants and their names: a concise dictionary. Oxford: Oxford University Press. ISBN 978-0-19-866189-4.

[26] The Complete Guide to Edible Wild Plants. United States Department of the Army. New York: Skyhorse Publishing. 2009. p. 64. ISBN 978-1-60239-692-0. OCLC 277203364.{{cite book}}: CS1 maint: others (link)

[27] "Weaving success: Manipur woman seeks to employ more locals in her lotus silk venture". The Print. PTI. 11 September 2022.

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@@ Line 1: / Line 1: @@
 {{Short description|Genus of aquatic flowering plants known as "lotus."}}
 {{Automatic taxobox
-| fossil_range = {{fossil range|Cretaceous|Recent}}
+| fossil_range = {{fossil range|Albian|recent|[[Albian]] to present<ref name="Li-2014"/>}}
 | image = Nelumno nucifera open flower - botanic garden adelaide2.jpg
-| image_caption = ''N. nucifera'' (sacred lotus)
+| image_caption = ''[[Nelumbo nucifera]]'' (sacred lotus)
+| image2 = American Lotus (Nelumbo lutea) 01.jpg
+| image2_caption = ''[[Nelumbo lutea]]'' (American lotus)
 | taxon = Nelumbo
 | authority = [[Michel Adanson|Adans.]]
 | subdivision_ranks = Species
-| subdivision =
+| subdivision = *''[[Nelumbo lutea]]''
+*''[[Nelumbo nucifera]]''
 *†''[[Nelumbo aureavallis]]''
 *†''[[Nelumbo changchangensis]]''
-*''[[Nelumbo lutea]]''
 *†''[[Nelumbo minima]]''
 *†''[[Nelumbo nipponica]]''
-*''[[Nelumbo nucifera]]''
 *†''[[Nelumbo orientalis]]''
 *†''[[Nelumbo protolutea]]''
+* 30+ more from fossil remains
 }}
-'''''Nelumbo''''' is a [[genus]] of [[aquatic plant]]s with large, showy [[flower]]s. Members are commonly called '''lotus''', though "[[List of plants known as lotus|lotus]]" is a name also applied to various other plants and plant groups, including the unrelated genus ''[[Lotus (genus)|Lotus]]''.  Members outwardly resemble those in the [[Family (biology)|family]] [[Nymphaeaceae]] ("water lilies"), but ''Nelumbo'' is actually very distant to Nymphaeaceae.  "Nelumbo" is derived from the [[Sinhala language|Sinhala]] word: {{lang-si|නෙළුම්}} ''neḷum'', the name for the lotus ''[[Nelumbo nucifera]]''.<ref name=HyamPank95/>
+'''''Nelumbo''''' {{IPAc-en|n|ɪ|ˈ|l|ʌ|m|b|oʊ}}<ref>{{Cite OED|nelumbo}}</ref> is a [[genus]] of [[aquatic plant]]s with large, showy [[flower]]s. Members are commonly called '''lotus''', though the name is also applied to [[List of plants known as lotus|various other plants and plant groups]], including the unrelated genus ''[[Lotus (genus)|Lotus]]''.  Members outwardly resemble those in the family [[Nymphaeaceae]] ("water lilies"), but ''Nelumbo'' is actually very distant from that family.
-There are only two known living species of lotus; ''N. nucifera'' is native to [[East Asia]], [[South Asia]] and [[Southeast Asia]] and is better-known. It is commonly cultivated; it is eaten and used in [[traditional Chinese medicine]]. This species is the [[floral emblem]] of both [[India]] and [[Vietnam]].
+''Nelumbo'' is an ancient genus, with dozens of species known from fossil remains since the [[Early Cretaceous]]. However, there are only two known living species of lotus. One is the better-known ''[[Nelumbo nucifera]]'', which is native to [[East Asia]], [[South Asia]], [[Southeast Asia]], and probably [[Australia]] and is commonly cultivated for consumption and use in [[traditional Chinese medicine]]. The other lotus is ''[[Nelumbo lutea]]'', which is [[Native species|native]] to [[North America]] and the [[Caribbean]]. Horticultural hybrids have been produced between these two [[allopatric speciation|allopatric species]].
+== Description ==
-The other lotus is ''[[Nelumbo lutea]]'' and is [[native plant|native]] to [[North America]] and the [[Caribbean]]. Horticultural hybrids have been produced between these two [[allopatric speciation|allopatric species]].
+===Ultrahydrophobicity===
-There are several fossil species known from [[Cretaceous]], [[Paleogene]] and [[Neogene]] aged strata throughout [[Eurasia]] and [[North America]].
+[[File:Nelumbo nucifera (Indian Lotus)- water drops W IMG 8657.jpg|thumb|left|Foliage of ''[[Nelumbo nucifera|N.&nbsp;nucifera]]'': an example of the [[lotus effect]] after rain]]
+The [[leaves]] of ''Nelumbo'' are highly water-repellent (i.e. they exhibit [[ultrahydrophobicity]]) and have given the name to what is called the [[lotus effect]].<ref>{{cite journal |last1=Darmanin |first1=Thierry |last2=Guittard |first2=Frédéric |name-list-style=vanc |date=1 June 2015 |title=Superhydrophobic and superoleophobic properties in nature |journal=Materials Today |volume=18 |issue=5 |pages=273–285 |doi=10.1016/j.mattod.2015.01.001 |doi-access=free}}</ref> Ultrahydrophobicity involves two criteria: a very high water contact angle between the droplet of water and the leaf surface, and a very low roll-off angle.<ref name="Marmur_2004">{{Cite journal |last=Marmur |first=Abraham |name-list-style=vanc |date=2004-04-01 |title=The Lotus Effect: Superhydrophobicity and Metastability |journal=Langmuir |volume=20 |issue=9 |pages=3517–3519 |doi=10.1021/la036369u |pmid=15875376}}</ref> This means that the water must contact the leaf surface at exactly one, minuscule point, and any manipulation of the leaf by changing its angle will result in the water droplet rolling off of the leaf.<ref name="Marmur_2004" /> Ultrahydrophobicity is conferred by the usually dense layer of papillae on the surface of the ''Nelumbo'' leaves, and the small, robust, waxy tubules that protrude off each papilla.<ref name="Zhang_2012">{{cite journal |last1=Zhang |first1=Youfa |last2=Wu |first2=Hao |last3=Yu |first3=Xinquan |last4=Chen |first4=Feng |last5=Wu |first5=Jie |name-list-style=vanc |date=March 2012 |title=Microscopic Observations of the Lotus Leaf for Explaining the Outstanding Mechanical Properties |journal=Journal of Bionic Engineering |volume=9 |issue=1 |pages=84–90 |doi=10.1016/S1672-6529(11)60100-5|s2cid=137076244 }}</ref> This helps reduce the area of contact between the water droplet and the leaf.<ref name="Zhang_2012" />
-==Species==
-[[File:American Lotus (Nelumbo lutea) 01.jpg|thumb|''[[Nelumbo lutea|N. lutea]]'' (American lotus)]]
-[[File:Lotus Nelumbo 'Mrs. Perry D. Slocum' Dried Seed Head 2000px.jpg|thumb|Nelumbo 'Mrs. Perry D. Slocum'- Dried seed pod]]
+Ultrahydrophobicity is said to confer a very important [[evolution]]ary advantage. As an aquatic plant with leaves that rest on the water's surface, the genus ''Nelumbo'' is characterized by its concentration of [[stoma]]ta on the upper [[Epidermis (botany)|epidermis]] of its leaves, unlike most other plants which concentrate their stomata on the lower [[Epidermis (botany)|epidermis]], underneath the leaf.<ref name="Zhang_2012" /> The collection of water on the upper epidermis, whether that be by rain, mist, or the nearby disturbance of water, is very detrimental to the leaf's ability to perform gas exchange through its stomata. Thus, ''Nelumbo's'' ultrahydrophobicity allows the water droplets to accumulate together very quickly, and then roll off of the leaf very easily at the slightest disturbance of the leaf, a process which allows its stomata to function normally without restriction due to blockage by water droplets.<ref>{{cite journal |vauthors=Ensikat HJ, Ditsche-Kuru P, Neinhuis C, Barthlott W |date=2011-03-10 |title=Superhydrophobicity in perfection: the outstanding properties of the lotus leaf |journal=Beilstein Journal of Nanotechnology |volume=2 |pages=152–61 |doi=10.3762/bjnano.2.19 |pmc=3148040 |pmid=21977427}}</ref>
-===Extant species===
-*''[[Nelumbo lutea]]'' <small>[[Carl Ludwig Willdenow|Willd.]]</small> &ndash; American lotus ([[Eastern United States]], [[Mexico]], [[Greater Antilles]], [[Honduras]])
-*''[[Nelumbo nucifera]]'' <small>[[Joseph Gaertner|Gaertn.]]</small> &ndash; sacred or Indian lotus, also known as the Rose of India and the sacred water lily of [[Hinduism]] and [[Buddhism]].<ref name=Kew>{{cite web |title=Nelumbo nucifera (sacred lotus) |url=http://www.kew.org/science-conservation/plants-fungi/nelumbo-nucifera-sacred-lotus | archive-url = https://web.archive.org/web/20140530010810/https://www.kew.org/science-conservation/plants-fungi/nelumbo-nucifera-sacred-lotus | archive-date = 30 May 2014 |publisher=Kew Royal Botanic Gardens |access-date=26 July 2015}}</ref> It is the [[Floral emblem|national flower]] of [[India]] and [[Vietnam]]. Its roots and [[Lotus seed|seeds]] are also used widely in cooking in East Asia, South Asia and Southeast Asia.
-[[File:Nelumbo nucifera LOTUS bud.jpg|thumb|''[[Nelumbo nucifera]]'' bud]]
-[[File:Lotus2mq.jpg|thumb|Microscopic water droplets resting above the leaf surface, allowing gas exchange to continue.]]
-===Fossil species===
+===Thermoregulation===
+An uncommon property of the genus ''Nelumbo'' is that it can [[Thermoregulation#In plants|generate heat]],<ref name="Watling_2006">{{cite journal |vauthors=Watling JR, Robinson SA, Seymour RS |date=April 2006 |title=Contribution of the alternative pathway to respiration during thermogenesis in flowers of the sacred lotus |journal=Plant Physiology |volume=140 |issue=4 |pages=1367–73 |doi=10.1104/pp.105.075523 |pmc=1435819 |pmid=16461386}}</ref> which it does by using the alternative oxidase pathway (AOX).<ref>{{cite journal |vauthors=Seymour RS, Schultze-Motel P, Lamprecht I |date=1 July 1998 |title=Heat production by sacred lotus flowers depends on ambient temperature, not light cycle |journal=Journal of Experimental Botany |volume=49 |issue=324 |pages=1213–1217 |doi=10.1093/jxb/49.324.1213 |doi-access=free}}</ref><ref name="Grant_2010">{{Cite journal |last1=Grant |first1=Nicole M. |last2=Miller |first2=Rebecca A. |last3=Watling |first3=Jennifer R. |last4=Robinson |first4=Sharon A. |name-list-style=vanc |date=2010-11-12 |title=Distribution of thermogenic activity in floral tissues of Nelumbo nucifera |journal=Functional Plant Biology |volume=37 |issue=11 |pages=1085–1095 |doi=10.1071/FP10024 |issn=1445-4416|url=https://ro.uow.edu.au/scipapers/5155 }}</ref> This pathway involves a different, alternative exchange of [[electron]]s from the usual pathway that electrons follow when generating energy in [[Mitochondrion|mitochondria]], known as the AOX, or alternative oxidase pathway.
-*†''[[Nelumbo aureavallis]]'' <small>[[Leo J. Hickey|Hickey]]</small> &ndash; [[Ypresian|Eocene]] ([[North Dakota]]), described from leaves found in the Golden Valley Formation in North Dakota, USA.<ref name="Hickey1977">{{cite book |title=Stratigraphy and Paleobotany of the Golden Valley Formation (Early Tertiary) of Western North Dakota |last=Hickey |first=Leo |name-list-style=vanc |year=1977 |publisher=Geological Society of America |location=Boulder, Colorado |isbn=978-0-8137-1150-8 |pages=[https://archive.org/details/stratigraphypale0000hick/page/110 110 & Plate 5] |url=https://archive.org/details/stratigraphypale0000hick/page/110 }}</ref>
-*†''[[Nelumbo changchangensis]]'' Eocene, ([[Hainan Island]], China), described from several fossils of leaves, seedpods, and rhizomes from the Eocene-aged strata in the Changchang Basin, of Hainan Island.
-*†''[[Nelumbo minima]]''  [[Pliocene]] ([[Netherlands]]), described from leaves and seedpods that suggest a very small plant.  Originally described as a member of the genus ''[[Nelumbites]]'', as "''Nelumbites minimus''."
-*†''[[Nelumbo nipponica]]'' [[Eocene]]-[[Miocene]], fossil leaves are known from Eocene-aged strata in [[Japan]], and Miocene-aged strata in [[Russia]].
-*†''[[Nelumbo orientalis]]'' [[Cretaceous]] (Japan), one of the oldest known species, fossils are found in Cretaceous-aged strata of Japan.
-*†''[[Nelumbo protolutea]]'' Eocene ([[Mississippi]]), fossils of leaves strongly suggest a plant similar in form to the American lotus.
+The typical pathway in plant mitochondria involves [[cytochrome]] complexes. The pathway used to generate heat in ''Nelumbo'' involves [[cyanide]]-resistant alternative oxidase, which is a different electron acceptor than the usual cytochrome complexes.<ref name="Hiroma_2011">{{Cite journal |last1=Hiroma |first1=Tatsuo |last2=Ito |first2=Kikukatsu |last3=Hara |first3=Michihiro |last4=Torisu |first4=Ryo |name-list-style=vanc |date=2011-06-01 |title=Analysis of the Lotus Thermoregulation System from the Perspective of Control Engineering |journal=Shokubutsu Kankyo Kogaku |language=ja |volume=23 |issue=2 |pages=52–58 |doi=10.2525/shita.23.52 |issn=1880-2028 |doi-access=free}}</ref> The plant also reduces [[ubiquitin]] concentrations while in [[thermogenesis]], which allows the AOX in the plant to function without degradation.<ref name="Wang_2015">{{cite journal |last1=Wang |first1=Ruohan |last2=Zhang |first2=Zhixiang |name-list-style=vanc |date=March 2015 |title=Floral thermogenesis: An adaptive strategy of pollination biology in Magnoliaceae |journal=Communicative & Integrative Biology |volume=8 |issue=1 |pages=e992746 |doi=10.4161/19420889.2014.992746 |pmc=4594551 |pmid=26844867}}</ref> Thermogenesis is restricted to the [[Receptacle (botany)|receptacle]], [[stamen]], and [[petal]]s of the flower, but each of these parts produce heat independently without relying on the heat production in other parts of the flower.<ref name="Li_2009">{{cite journal |vauthors=Li JK, Huang SQ |date=May 2009 |title=Flower thermoregulation facilitates fertilization in Asian sacred lotus |journal=Annals of Botany |volume=103 |issue=7 |pages=1159–63 |doi=10.1093/aob/mcp051 |pmc=2707905 |pmid=19282320}}</ref>
-==Classification==
-There is residual disagreement over which family the genus should be placed in.  Traditional classification systems recognized ''Nelumbo'' as part of the Nymphaeaceae, but traditional taxonomists were likely misled by [[convergent evolution]] associated with an evolutionary shift from a terrestrial to an aquatic lifestyle.  In the older classification systems it was recognized under the [[Order (biology)|biological order]] [[Nymphaeales]] or Nelumbonales. ''Nelumbo'' is currently recognized as the [[monotypic taxon|only living genus]] in [[Nelumbonaceae]], one of several distinctive families in the [[eudicot]] order of the [[Proteales]].  Its closest living relatives, the ([[Proteaceae]] and [[Platanaceae]]), are shrubs or trees.
+There are several theories about the function of thermogenesis, especially in an aquatic genus such as ''Nelumbo''. The most common theory posits that thermogenesis in flowers attracts [[pollinator]]s, for a variety of reasons. Heated flowers may attract insect pollinators. As the pollinators warm themselves while resting inside the flower, they deposit and pick up [[pollen]] onto and from the flower.<ref name="Watling_2006" /> The thermogenic environment might also be conducive to pollinator [[mating]]; pollinators may require a certain temperature for reproduction. By providing an ideal thermogenic environment, the flower is pollinated by mating pollinators.<ref name="Miller_2009">{{cite journal |vauthors=Miller RE, Watling JR, Robinson SA |year=2009 |title=Functional transition in the floral receptacle of the sacred lotus (Nelumbo nucifera): from thermogenesis to photosynthesis. |url=https://scholars.uow.edu.au/display/publication26931 |journal=Functional Plant Biology |volume=36 |issue=5 |pages=471–480 |doi=10.1071/FP08326|pmid=32688661 |s2cid=54588650 }}</ref> Others theorize that heat production facilitates the release of [[Volatile organic compound|volatile]] compounds into the air to attract pollinators flying over water, or that the heat is recognizable in the dark by thermo-sensitive pollinators. None have been conclusively proven to be more plausible than the others.<ref name="Wagner_2008">{{cite journal |vauthors=Wagner AM, Krab K, Wagner MJ, Moore AL |date=2008-07-01 |title=Regulation of thermogenesis in flowering Araceae: the role of the alternative oxidase |journal=Biochimica et Biophysica Acta (BBA) - Bioenergetics |volume=1777 |issue=7–8 |pages=993–1000 |doi=10.1016/j.bbabio.2008.04.001 |pmid=18440298 }}</ref>
-The [[leaf|leaves]] of ''Nelumbo'' can be distinguished from those of genera in the Nymphaeaceae as they are [[Glossary of leaf morphology|peltate]], that is they have fully circular leaves. ''[[Nymphaea]]'', on the other hand, has a single characteristic notch from the edge in to the center of the lily pad. The [[fruit|seedpod]] of ''Nelumbo'' is very distinctive.
+After [[anthesis]], the receptacle of the lotus transitions from a primarily thermogenic to a [[Photosynthesis|photosynthetic]] structure, as seen in the rapid and dramatic increase in [[photosystem]]s, photosynthetically involved pigments, electron transport rates, and the presence of [[Carbon-13|13<sup href="Nelumbo lutea">C</sup>]] in the receptacle and petals, all of which assist in increasing photosynthesis rates.<ref name="Grant_2008">{{cite journal |vauthors=Grant NM, Miller RE, Watling JR, Robinson SA |date=2008-02-01 |title=Synchronicity of thermogenic activity, alternative pathway respiratory flux, AOX protein content, and carbohydrates in receptacle tissues of sacred lotus during floral development |journal=Journal of Experimental Botany |volume=59 |issue=3 |pages=705–14 |doi=10.1093/jxb/erm333 |pmid=18252702 |doi-access=free|hdl=2440/52460 |hdl-access=free }}</ref> After this transition, all thermogenesis in the flower is lost. Pollinators do not need to be attracted once the [[Ovary (botany)|ovary]] is fertilized, and thus the receptacle's resources are better used when it is photosynthesizing to produce [[carbohydrate]]s that can increase plant biomass or fruit mass.<ref name="Miller_2009" />
-===APG===
-[[File:Nelumbo nucifera (Indian Lotus)- water drops W IMG 8657.jpg|thumb|Foliage of ''Nelumbo nucifera'': an example of the [[lotus effect]] after rain.]]
-The [[APG IV system]] of 2016, recognizes Nelumbonaceae as a distinct family and places it in the order Proteales in the eudicot [[clade]], as do the earlier [[APG III system|APG III]] and [[APG II system]]s.<ref name=APGIV/>
+Other plants utilize thermoregulation in their life cycles. Among these is the [[Symplocarpus foetidus|eastern skunk cabbage]], which heats itself to melt any ice above it, and push through the ground in early spring.<ref>{{Cite journal |last1=Ito |first1=Kikukatsu |last2=Ito |first2=Takanori |last3=Onda |first3=Yoshihiko |last4=Uemura |first4=Matsuo |name-list-style=vanc |date=2004-03-15 |title=Temperature-Triggered Periodical Thermogenic Oscillations in Skunk Cabbage (Symplocarpus foetidus) |journal=Plant and Cell Physiology |volume=45 |issue=3 |pages=257–264 |doi=10.1093/pcp/pch038 |issn=0032-0781 |pmid=15047873 |doi-access=free}}</ref> Also, the [[Konjac|elephant yam]], which heats its flowers to attract pollinators. In addition, the [[carrion flower]], which heats itself to disperse water vapor through the air, carrying its scent further, thus attracting more pollinators.
-===Earlier classification systems===
-The [[Cronquist system]] of 1981 recognizes the family but places it in the water lily order [[Nymphaeales]].  The [[Dahlgren system]] of 1985 and [[Thorne system]] of 1992 both recognize the family and place it in its own order, Nelumbonales.  The USDA still classifies the lotus family within the water lily order.<ref>http://plants.sc.egov.usda.gov/core/profile?symbol=NENU2, Click on Classification tab, three to the right of the General tab under which the webpage opens. (It is all the same web address.)</ref>
+[[File:Nelumbo nucifera LOTUS bud.jpg|thumb|upright=.8|''[[Nelumbo nucifera]]'' bud]]
-==Characteristics==
-===Ultrahydrophobicity===
+=== Similar species ===
+The leaves of ''Nelumbo'' can be distinguished from those of genera in the family [[Nymphaeaceae]] as they are [[Glossary of leaf morphology|peltate]], that is they have fully circular leaves. ''[[Nymphaea]]'', on the other hand, has a single characteristic notch from the edge in to the center of the lily pad. The [[fruit|seedpod]] of ''Nelumbo'' is very distinctive.
-The leaves of ''Nelumbo'' are highly water-repellent (i.e. they exhibit [[ultrahydrophobicity]]) and have given the name to what is called the [[lotus effect]].<ref>{{cite journal|last1=Darmanin|first1=Thierry|last2=Guittard|first2=Frédéric | name-list-style = vanc |title=Superhydrophobic and superoleophobic properties in nature|journal=Materials Today|date=1 June 2015|volume=18|issue=5|pages=273–285|doi=10.1016/j.mattod.2015.01.001 |doi-access=free}}</ref> Ultrahydrophobicity involves two criteria: a very high water contact angle between the droplet of water and the leaf surface, and a very low roll-off angle.<ref name="Marmur_2004">{{Cite journal|last=Marmur|first=Abraham | name-list-style = vanc |date=2004-04-01|title=The Lotus Effect: Superhydrophobicity and Metastability |journal=Langmuir|volume=20|issue=9|pages=3517–3519|doi=10.1021/la036369u|pmid=15875376 }}</ref> This means that the water must contact the leaf surface at exactly one, minuscule point, and any manipulation of the leaf by changing its angle will result in the water droplet rolling off of the leaf.<ref name="Marmur_2004" /> Ultrahydrophobicity is conferred by the usually dense layer of papillae on the surface of the ''Nelumbo'' leaves, and the small, robust, waxy tubules that protrude off each papillae.<ref name="Zhang_2012">{{cite journal | last1 = Zhang | first1 = Youfa | last2 = Wu | first2 = Hao | last3 = Yu | first3 = Xinquan | last4 = Chen | first4 = Feng | last5 = Wu | first5 = Jie | name-list-style = vanc | title = Microscopic Observations of the Lotus Leaf for Explaining the Outstanding Mechanical Properties | journal = Journal of Bionic Engineering | date = March 2012 | volume = 9 | issue = 1 | pages = 84–90 | doi = 10.1016/S1672-6529(11)60100-5 }}</ref> This helps reduce the area of contact between the water droplet and the leaf.<ref name="Zhang_2012" />
+== Taxonomy ==
-Ultrahydrophobicity is said to confer a very important [[evolution]]ary advantage. As an aquatic plant with leaves that rest on the water's surface, the genus ''Nelumbo'' is characterized by its concentration of [[stoma]]ta on the upper [[Epidermis (botany)|epidermis]] of its leaves, unlike most other plants which concentrate their stomata on the lower [[Epidermis (botany)|epidermis]], underneath the leaf.<ref name="Zhang_2012" /> The collection of water on the upper epidermis, whether that be by rain, mist, or the nearby disturbance of water, is very detrimental to the leaf's ability to perform gas exchange through its stomata. Thus, ''Nelumbo's'' ultrahydrophobicity allows the water droplets to accumulate together very quickly, and then roll off of the leaf very easily at the slightest disturbance of the leaf, a process which allows its stomata to function normally without restriction due to blockage by water droplets.<ref>{{cite journal | vauthors = Ensikat HJ, Ditsche-Kuru P, Neinhuis C, Barthlott W | title = Superhydrophobicity in perfection: the outstanding properties of the lotus leaf | journal = Beilstein Journal of Nanotechnology | volume = 2 | pages = 152–61 | date = 2011-03-10 | pmid = 21977427 | pmc = 3148040 | doi = 10.3762/bjnano.2.19 }}</ref>
-===Thermoregulation===
+=== Taxonomic history ===
+The [[Cronquist system]] of 1981 recognizes the family [[Nelumbonaceae]] but places it in the water lily order [[Nymphaeales]]. The [[Dahlgren system]] of 1985 and [[Thorne system]] of 1992 both recognize the family and place it in its own order, Nelumbonales. The [[United States Department of Agriculture]] still classifies the lotus family within the water lily order.<ref>{{Cite PLANTS |symbol=NENU2 |taxon=''Nelumbo nucifera'' |access-date=2024-11-03}}</ref>
-A unique property of the genus ''Nelumbo'' is that it can generate heat,<ref name="Watling_2006">{{cite journal | vauthors = Watling JR, Robinson SA, Seymour RS | title = Contribution of the alternative pathway to respiration during thermogenesis in flowers of the sacred lotus | journal = Plant Physiology | volume = 140 | issue = 4 | pages = 1367–73 | date = April 2006 | pmid = 16461386 | pmc = 1435819 | doi = 10.1104/pp.105.075523 }}</ref> which it does by using the alternative oxidase pathway (AOX).<ref>{{cite journal | vauthors = Seymour RS, Schultze-Motel P, Lamprecht I | title = Heat production by sacred lotus flowers depends on ambient temperature, not light cycle|journal=Journal of Experimental Botany|date=1 July 1998|volume=49|issue=324|pages=1213–1217|doi=10.1093/jxb/49.324.1213|doi-access=free}}</ref><ref name="Grant_2010">{{Cite journal|last1=Grant|first1=Nicole M.|last2=Miller|first2=Rebecca A.|last3=Watling|first3=Jennifer R.|last4=Robinson|first4=Sharon A.| name-list-style = vanc |date=2010-11-12|title=Distribution of thermogenic activity in floral tissues of Nelumbo nucifera |journal=Functional Plant Biology|volume=37|issue=11|pages=1085–1095|doi=10.1071/FP10024|issn=1445-4416}}</ref> This pathway involves a different, alternative exchange of [[electron]]s from the usual pathway that electrons follow when generating energy in [[Mitochondrion|mitochondria]], known as the AOX, or alternative oxidase pathway.
+There is residual disagreement over which family the genus should be placed in. Traditional classification systems recognized ''Nelumbo'' as part of the Nymphaeaceae, but traditional taxonomists were likely misled by [[convergent evolution]] associated with an evolutionary shift from a terrestrial to an aquatic lifestyle. In the older classification systems it was recognized under the order Nymphaeales or Nelumbonales.
-The typical pathway in plant mitochondria involves [[cytochrome]] complexes. The pathway used to generate heat in ''Nelumbo'' involves [[cyanide]]-resistant alternative oxidase, which is a different electron acceptor than the usual cytochrome complexes.<ref name="Hiroma_2011">{{Cite journal|last1=Hiroma |first1=Tatsuo|last2= Ito |first2=Kikukatsu|last3= Hara |first3=Michihiro|last4= Torisu |first4=Ryo| name-list-style = vanc |date=2011-06-01|title=Analysis of the Lotus Thermoregulation System from the Perspective of Control Engineering|journal=Shokubutsu Kankyo Kogaku|language=ja|volume=23|issue=2|pages=52–58|doi=10.2525/shita.23.52|issn=1880-2028|doi-access=free}}</ref> The plant also reduces [[ubiquitin]] concentrations while in [[thermogenesis]], which allows the AOX in the plant to function without degradation<ref name="Wang_2015">{{cite journal|last1=Wang|first1=Ruohan|last2=Zhang|first2=Zhixiang | name-list-style = vanc |title=Floral thermogenesis: An adaptive strategy of pollination biology in Magnoliaceae|journal=Communicative & Integrative Biology|date= March 2015 | volume = 8 | issue = 1 | pages = e992746 | doi = 10.4161/19420889.2014.992746 |pmid=26844867|pmc=4594551}}</ref> Thermogenesis is restricted to the [[Receptacle (botany)|receptacle]], [[stamen]], and [[petal]]s of the flower, but each of these parts produce heat independently without relying on the heat production in other parts of the flower.<ref name="Li_2009">{{cite journal | vauthors = Li JK, Huang SQ | title = Flower thermoregulation facilitates fertilization in Asian sacred lotus | journal = Annals of Botany | volume = 103 | issue = 7 | pages = 1159–63 | date = May 2009 | pmid = 19282320 | pmc = 2707905 | doi = 10.1093/aob/mcp051 }}</ref>
+=== Modern classification ===
-There are several theories about the function of thermogenesis, especially in an aquatic genus such as ''Nelumbo''. The most common theory posits that thermogenesis in flowers attracts [[pollinator]]s, for a variety of reasons. Heated flowers may attract insect pollinators. As the pollinators warm themselves while resting inside the flower, they deposit and pick up [[pollen]] onto and from the flower.<ref name="Watling_2006" /> The thermogenic environment might also be conducive to pollinator [[mating]] - pollinators may require a certain temperature for reproduction. By providing an ideal thermogenic environment, the flower is pollinated by mating pollinators.<ref name = "Miller_2009">{{cite journal | vauthors = Miller RE, Watling JR, Robinson SA | year = 2009 | title = Functional transition in the floral receptacle of the sacred lotus (Nelumbo nucifera): from thermogenesis to photosynthesis. | journal = Functional Plant Biology | volume = 36 | issue = 5 | pages = 471–480 | url = https://scholars.uow.edu.au/display/publication26931 | doi = 10.1071/FP08326 }}</ref> Others theorize that heat production facilitates the release of [[Volatile organic compound|volatile]] compounds into the air to attract pollinators flying over water, or that the heat is recognizable in the dark by thermo-sensitive pollinators. None have been conclusively proven to be more plausible than the others.<ref name="Wagner_2008">{{cite journal | vauthors = Wagner AM, Krab K, Wagner MJ, Moore AL | title = Regulation of thermogenesis in flowering Araceae: the role of the alternative oxidase | journal = Biochimica et Biophysica Acta (BBA) - Bioenergetics | volume = 1777 | issue = 7–8 | pages = 993–1000 | date = 2008-07-01 | pmid = 18440298 | doi = 10.1016/j.bbabio.2008.04.001 | doi-access = free }}</ref>
+''Nelumbo'' is currently recognized as the [[monotypic taxon|only living genus]] in Nelumbonaceae, one of several distinctive families in the [[eudicot]] order of the [[Proteales]]. Its closest living relatives, the ([[Proteaceae]] and [[Platanaceae]]), are shrubs or trees.
+The [[APG IV system]] of 2016 recognizes Nelumbonaceae as a distinct family and places it in the order Proteales in the eudicot [[clade]], as do the earlier [[APG III system|APG III]] and [[APG II system]]s.<ref name="APGIV" />
+=== Phylogeny ===
+There are several fossil species known from [[Cretaceous]], [[Paleogene]] and [[Neogene]] aged strata throughout [[Eurasia]] and [[North America]]. Despite the ancient origins of this genus and the wide geographic separation of the two extant species (''N. nucifera'' and ''N. lutea''), phylogenetic evidence indicates that they diverged rather recently, during the early [[Pleistocene]] (about 2 million years ago).<ref>{{Cite journal |last1=Wu |first1=Zhihua |last2=Gui |first2=Songtao |last3=Quan |first3=Zhiwu |last4=Pan |first4=Lei |last5=Wang |first5=Shuzhen |last6=Ke |first6=Weidong |last7=Liang |first7=Dequan |last8=Ding |first8=Yi |date=2014-11-19 |title=A precise chloroplast genome of Nelumbo nucifera (Nelumbonaceae) evaluated with Sanger, Illumina MiSeq, and PacBio RS II sequencing platforms: insight into the plastid evolution of basal eudicots |journal=BMC Plant Biology |volume=14 |issue=1 |page=289 |doi=10.1186/s12870-014-0289-0 |issn=1471-2229 |pmc=4245832 |pmid=25407166 |doi-access=free }}</ref>
+=== Species ===
+[[File:Lotus Nelumbo 'Mrs. Perry D. Slocum' Dried Seed Head 2000px.jpg|thumb|Dried seed pod from ''Nelumbo'' 'Mrs. Perry D. Slocum', a cross between the two extant species]]
+==== Extant species ====
+*''[[Nelumbo lutea]]'' <small>[[Carl Ludwig Willdenow|Willd.]]</small> &ndash; American lotus ([[Eastern United States]], [[Mexico]], [[Greater Antilles]], [[Honduras]])
+*''[[Nelumbo nucifera]]'' <small>[[Joseph Gaertner|Gaertn.]]</small> &ndash; sacred or Indian lotus, also known as the Rose of India and the sacred water lily of [[Hinduism]] and [[Buddhism]].<ref name="Kew">{{cite web |title=Nelumbo nucifera (sacred lotus) |url=http://www.kew.org/science-conservation/plants-fungi/nelumbo-nucifera-sacred-lotus | archive-url = https://web.archive.org/web/20140530010810/https://www.kew.org/science-conservation/plants-fungi/nelumbo-nucifera-sacred-lotus | archive-date = 30 May 2014 |publisher=Kew Royal Botanic Gardens |access-date=26 July 2015}}</ref> It is the [[Floral emblem|national flower]] of [[India]] and [[Vietnam]]. Its roots and [[Lotus seed|seeds]] are also used widely in cooking in East Asia, South Asia and Southeast Asia.
+==== Fossil species ====
+Nearly 30 fossil species are known from the mid-Cretaceous to the present.<ref name="Li-2014">{{Cite journal |last1=Li |first1=Ya |last2=Svetlana |first2=Popova |last3=Yao |first3=Jianxin |last4=Li |first4=Chengsen |date=2014 |title=A Review on the Taxonomic, Evolutionary and Phytogeographic Studies of the Lotus Plant (Nelumbonaceae: Nelumbo ) |url=https://onlinelibrary.wiley.com/doi/10.1111/1755-6724.12287 |journal=Acta Geologica Sinica - English Edition |language=en |volume=88 |issue=4 |pages=1252–1261 |doi=10.1111/1755-6724.12287 |bibcode=2014AcGlS..88.1252L |issn=1000-9515}}</ref>
+*†''[[Nelumbo aureavallis]]'' <small>[[Leo J. Hickey|Hickey]]</small> &ndash; [[Ypresian|Eocene]] ([[North Dakota]]), described from leaves found in the Golden Valley Formation in North Dakota, USA.<ref name="Hickey1977">{{cite book |title=Stratigraphy and Paleobotany of the Golden Valley Formation (Early Tertiary) of Western North Dakota |last=Hickey |first=Leo |name-list-style=vanc |year=1977 |publisher=Geological Society of America |location=Boulder, Colorado |isbn=978-0-8137-1150-8 |pages=[https://archive.org/details/stratigraphypale0000hick/page/110 110 & Plate 5] |url=https://archive.org/details/stratigraphypale0000hick/page/110 }}</ref>
+*†''[[Nelumbo changchangensis]]'' Eocene, ([[Hainan Island]], China), described from several fossils of leaves, seedpods, and rhizomes from the Eocene-aged strata in the Changchang Basin, of Hainan Island.
+*†''[[Nelumbo choffati]]'' [[Early Cretaceous]] ([[Portugal]]), leaves known from the [[Albian]]. One of the earliest known species.<ref name="Li-2014" />
+*†''[[Nelumbo jiayinensis]]'' [[Late Cretaceous]], ([[Heilongjiang]], China), leaves described from the [[Santonian]]-aged [[Yong'ancun Formation]]<ref>{{Cite journal |last1=Liang |first1=Fei |last2=Sun |first2=Ge |last3=Yang |first3=Tao |last4=Bai |first4=Shuchong |date=April 2018 |title=Nelumbo jiayinensis sp. nov. from the Upper Cretaceous Yong'ancun Formation in Jiayin, Heilongjiang, Northeast China |url=https://doi.org/10.1016/j.cretres.2017.11.007 |journal=Cretaceous Research |volume=84 |pages=134–140 |doi=10.1016/j.cretres.2017.11.007 |bibcode=2018CrRes..84..134L |issn=0195-6671}}</ref>
+*†''[[Nelumbo lusitanica]]'' Early Cretaceous (Portugal), leaves known from the Albian. One of the earliest known species.<ref name="Li-2014" />
+*†''[[Nelumbo minima]]''  [[Pliocene]] ([[Netherlands]]), described from leaves and seedpods that suggest a very small plant.  Originally described as a member of the genus ''[[Nelumbites]]'', as "''Nelumbites minimus''."
+*†''[[Nelumbo nipponica]]'' [[Eocene]]-[[Miocene]], fossil leaves are known from Eocene-aged strata in [[Japan]], and Miocene-aged strata in [[Russia]].
+*†''[[Nelumbo orientalis]]'' Cretaceous (Japan), fossils found in Cretaceous-aged strata of Japan. The [[Sarao Formation]], which they are known from, was formerly considered of Early Cretaceous age, but more recent studies support a [[Maastrichtian]] age for it.<ref>{{Cite web |title=Discovery of Nelumbo from the Asuwa Flora (Upper Cretaceous) in Fukui Prefecture in the Inner Side of Central Japan |url=https://gbank.gsj.jp/geolis/geolis_link/88800443/en |access-date=2024-07-19 |website=gbank.gsj.jp}}</ref><ref>{{Cite journal |last1=佐沢 |first1=豪 |last2=ルグラン |first2=ジュリアン |last3=矢部 |first3=淳 |last4=上松 |first4=佐知子 |last5=指田 |first5=勝男 |date=2020 |title=福井県東部，池田町に分布する上部白亜系足羽層の年代-花粉層位学的アプローチ |url=https://www.jstage.jst.go.jp/article/geosoc/126/4/126_2020.0004/_article/-char/ja/ |journal=地質学雑誌 |volume=126 |issue=4 |pages=215–221 |doi=10.5575/geosoc.2020.0004|doi-access=free }}</ref>
+*†''[[Nelumbo protolutea]]'' Eocene ([[Mississippi]]), fossils of leaves strongly suggest a plant similar in form to the American lotus.
+*†''[[Nelumbo weymouthi]]'' Early Cretaceous ([[Wyoming]], US), leaves known from the Albian. One of the earliest known species.<ref name="Li-2014" />
+=== Etymology ===
-After [[anthesis]], the receptacle of the lotus transitions from a primarily thermogenic to a [[Photosynthesis|photosynthetic]] structure, as seen in the rapid and dramatic increase in [[photosystem]]s, photosynthetically involved pigments, electron transport rates, and the presence of [[Carbon-13|13<sup href="Nelumbo lutea">C</sup>]] in the receptacle and petals, all of which assist in increasing photosynthesis rates.<ref name="Grant_2008">{{cite journal | vauthors = Grant NM, Miller RE, Watling JR, Robinson SA | title = Synchronicity of thermogenic activity, alternative pathway respiratory flux, AOX protein content, and carbohydrates in receptacle tissues of sacred lotus during floral development | journal = Journal of Experimental Botany | volume = 59 | issue = 3 | pages = 705–14 | date = 2008-02-01 | pmid = 18252702 | doi = 10.1093/jxb/erm333 | doi-access = free }}</ref> After this transition, all thermogenesis in the flower is lost. Pollinators do not need to be attracted once the [[Ovary (botany)|ovary]] is fertilized, and thus the receptacle's resources are better used when it is photosynthesizing to produce [[carbohydrate]]s that can increase plant biomass or fruit mass.<ref name="Miller_2009" />
+The genus name is derived from {{langx|si|නෙළුම්}} ''neḷum'', the name for ''Nelumbo nucifera''.<ref name="HyamPank95" />
+==Uses==
-Other plants utilize thermoregulation in their life cycles. Among these is the [[Symplocarpus foetidus|eastern skunk cabbage]], which heats itself to melt any ice above it, and push through the ground in early spring,.<ref>{{Cite journal|last1=Ito|first1=Kikukatsu|last2=Ito|first2=Takanori|last3=Onda|first3=Yoshihiko|last4=Uemura|first4=Matsuo| name-list-style = vanc |date=2004-03-15|title=Temperature-Triggered Periodical Thermogenic Oscillations in Skunk Cabbage (Symplocarpus foetidus) |journal=Plant and Cell Physiology|volume=45|issue=3|pages=257–264|doi=10.1093/pcp/pch038|pmid=15047873|issn=0032-0781|doi-access=free}}</ref> Also, the [[Konjac|elephant yam]], which heats its flowers to attract pollinators. In addition, the [[carrion flower]], which heats itself to disperse water vapor through the air, carrying its scent further, thus attracting more pollinators.
+ [[File:JaRenkonLotus14R.jpg|thumb|Vinegared lotus root slices with ginger and citron as eaten in Japan]] The entire plant can be eaten either raw or cooked. The underwater portion is high in [[starch]]. The fleshy parts can be dug from the mud and baked or boiled. The young leaves can be boiled. The seeds are palatable and can be eaten raw or dried and ground into flour.<ref>{{Cite book |title=The Complete Guide to Edible Wild Plants |publisher=[[Skyhorse Publishing]] |others=[[United States Department of the Army]] |year=2009 |isbn=978-1-60239-692-0 |location=New York |page=64 |language=en-US |oclc=277203364}}</ref> The stem fibers are also used to make [[lotus silk]].<ref>{{cite news |url=https://theprint.in/india/weaving-success-manipur-woman-seeks-to-employ-more-locals-in-her-lotus-silk-venture/1124204/ |title=Weaving success: Manipur woman seeks to employ more locals in her lotus silk venture |agency=[[Press Trust of India|PTI]] |work=[[The Print]] |date=11 September 2022}}</ref>
+==Culture==
-==Cultural significance==
-{{main|Nelumbo nucifera}}
+{{Main|Nelumbo nucifera}}
-The sacred lotus, ''N. nucifera'', is sacred in both [[Hindu iconography|Hinduism]] and [[Buddhist symbolism|Buddhism]].<ref name=Kew/>
+The sacred lotus, ''N. nucifera'', is sacred in both [[Hindu iconography|Hinduism]] and [[Buddhist symbolism|Buddhism]].<ref name=Kew/> It is the [[floral emblem]] of both [[India]] and [[Vietnam]].
+{{-}}
 == References ==
 {{Reflist|refs=
-<ref name=HyamPank95>{{Cite book | vauthors = Hyam R, Pankhurst RJ |year=1995 |title=Plants and their names : a concise dictionary |location=Oxford |publisher=Oxford University Press |isbn=978-0-19-866189-4 }}</ref>
+<ref name=HyamPank95>{{Cite book | vauthors = Hyam R, Pankhurst RJ |year=1995 |title=Plants and their names: a concise dictionary |location=Oxford |publisher=Oxford University Press |isbn=978-0-19-866189-4 }}</ref>
-<ref name=APGIV>{{Cite journal|authors=Angiosperm Phylogeny Group|year=2016|title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV|journal=Botanical Journal of the Linnean Society|volume=181|issue=1|pages=1–20|issn=0024-4074|doi=10.1111/boj.12385|doi-access=free}}</ref>
+<ref name=APGIV>{{Cite journal|author=Angiosperm Phylogeny Group |year=2016|title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV|journal=Botanical Journal of the Linnean Society|volume=181|issue=1|pages=1–20|issn=0024-4074|doi=10.1111/boj.12385|doi-access=free}}</ref>
 }}
@@ Line 90: / Line 107: @@
 {{Wikispecies}}
-* [http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Nelumbonaceae links at CSDL]
+* [http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Nelumbonaceae links at CSDL] {{Webarchive|url=https://web.archive.org/web/20081012212023/http://www.csdl.tamu.edu/FLORA/cgi/gateway_family?fam=Nelumbonaceae |date=2008-10-12 }}
 {{Taxonbar|from=Q207427}}
+{{Authority control}}
 [[Category:Nelumbo| ]]
 [[Category:Eudicot genera]]
+[[Category:Extant Albian first appearances]]