Biology:Gymnosperm

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The gymnosperms (/ˈdʒɪmnəˌspɜːrmz, -noʊ-/ (About this soundlisten) nə-spurmz, -⁠noh-; from Ancient Greek γυμνός, gymnós 'naked' and σπέρμα, spérma 'seed', thus 'naked seed') are a group of woody, perennial seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include conifers, cycads, ginkgos, and gnetophytes, forming the clade Gymnospermae.[1] The name is based on the unenclosed condition of their seeds (called ovules in their unfertilized state). The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo.

The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase.[2] The term "gymnosperm" is often used in paleobotany to refer to (the paraphyletic group of) all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.[3]

The gymnosperms and angiosperms together constitute the spermatophytes or seed plants. The spermatophytes are subdivided into five divisions, the angiosperms and four divisions of gymnosperms: the Cycadophyta, Ginkgophyta, Gnetophyta, and Pinophyta (also known as Coniferophyta). Newer classification place the gnetophytes among the conifers.[4] Numerous extinct seed plant groups are recognised including those considered pteridosperms/seed ferns, as well other groups like the Bennettitales.[5]

By far the largest group of living gymnosperms are the conifers (pines, cypresses, and relatives), followed by cycads, gnetophytes (Gnetum, Ephedra and Welwitschia), and Ginkgo biloba (a single living species). About 65% of gymnosperms are dioecious,[6] but conifers are almost all monoecious.[7] Some genera have ectomycorrhiza fungal associations with roots (Pinus),[8] while in some others (Cycas) small specialised roots called coralloid roots are associated with nitrogen-fixing cyanobacteria.[9]

Diversity and origin

Encephalartos sclavoi cone

Over 1,000 living species of gymnosperm exist.[10] It was previously widely accepted that the gymnosperms originated in the Late Carboniferous period, replacing the lycopsid rainforests of the tropical region, but more recent phylogenetic evidence indicates that they diverged from the ancestors of angiosperms during the Early Carboniferous.[11][12] The radiation of gymnosperms during the late Carboniferous appears to have resulted from a whole genome duplication event around 319 million years ago.[13] Early characteristics of seed plants are evident in fossil progymnosperms of the late Devonian period around 383 million years ago. It has been suggested that during the mid-Mesozoic era, pollination of some extinct groups of gymnosperms was by extinct species of scorpionflies that had specialized proboscides for feeding on pollination drops. The scorpionflies likely engaged in pollination mutualisms with gymnosperms, long before the similar and independent coevolution of nectar-feeding insects on angiosperms.[14][15] Some mid-Mesozoic gymnosperms were pollinated by kalligrammatid lacewings.[16]

All gymnosperms are perennial woody plants.[17] Unlike in other extant gymnosperms the soft and highly parenchymatous wood in cycads is poorly lignified,[18] and their main structural support comes from an armor of sclerenchymatous leaf bases covering the stem,[19] with the exception of species with underground stems.[20] There are no herbaceous gymnosperms and compared to angiosperms they occupy fewer ecological niches, but the gymnosperms include parasites (Parasitaxus), epiphytes (Zamia pseudoparasitica) and rheophytes (Retrophyllum minus).[21]

Conifers are by far the most abundant extant group of gymnosperms with six to eight families, with a total of 65–70 genera and 600–630 species (696 accepted names).[22] Most conifers are evergreens.[23] Conifers produce their seeds inside a protective cone called a strobilus. Most species are monoecious, with male and female cones on the same tree.[24][25] All conifers are wind-pollinated.[26]

Cycads, small palm-like trees,[1] are the next most abundant group of gymnosperms, with two or three families, 11 genera, and approximately 338 species. A majority of cycads are native to tropical climates and are most abundantly found in regions near the equator. The other extant groups are the 95–100 species of Gnetophytes and one species of Ginkgo. The ginkgo or maidenhair trees are tall and have bilobed leaves, while gnetophytes are a diverse groups of plants and shrubs including the horizontally growing Welwitschia.[5]

Gymnosperms are the most threatened of all plant groups.[27]

Evolution

Phylogeny

The phylogeny of the gymnosperms has been revised by multiple authors.[28][29][30][31]

Gymnospermae
 
Ginkgoidae

Ginkgoaceae

 
Cycadidae
Cycadales
 

Cycadaceae

 
 

Zamiaceae

 
 
 
 
 
Pinidae
 
Gnetales
 

Ephedraceae

 
 
 

Gnetaceae

 
 

Welwitschiaceae

 
 
 
Pinales

Pinaceae

 
 
 
Cupressidae
Araucariales
 

Araucariaceae

 
 

Podocarpaceae

 
 
Cupressales
 

Sciadopityaceae

 
 
Taxaceae
 

Cephalotaxaceae

 
 

Taxaceae

 
 
 

Cupressaceae

 
 
 
 
 
 

Taxonomy

A formal classification of the living gymnosperms is the "Acrogymnospermae", which form a monophyletic group within the spermatophytes.[32][33] The wider "Gymnospermae" group includes extinct gymnosperms and is thought to be paraphyletic. The fossil record of gymnosperms includes many distinctive taxa that do not belong to the four modern groups, including seed-bearing trees that have a somewhat fern-like vegetative morphology (the so-called "seed ferns" or pteridosperms).[34] When fossil gymnosperms such as these and the Bennettitales, glossopterids, and Caytonia are considered, it is clear that angiosperms are nested within a larger gymnospermae clade, although which group of gymnosperms is their closest relative remains unclear.

The extant gymnosperms include 12 main families and 83 genera which contain more than 1000 known species.[10][33][35]

Subclass Cycadidae

Subclass Ginkgoidae

Subclass Gnetidae

Subclass Pinidae

Extinct groupings

Life cycle

A gymnosperm lifecycle

Gymnosperms, like all vascular plants, have a sporophyte-dominant life cycle, which means they spend most of their life cycle with diploid cells, while the gametophyte (gamete-bearing phase) is relatively short-lived. Like all seed plants, they are heterosporous, having two spore types, microspores (male) produced in microsporangium and megaspores (female) produced in megasporangium that are typically present in pollen cones or ovulate cones respectively. The microsporangium is carried by microsporophyll (modified leaf) and seeds are carried by ovuliferous scales in the male and female cones respectively.[1][36] The exception is the females in the cycad genus Cycas, which form a loose structure called megasporophylls instead of cones.[37] As with all heterosporous plants, the gametophytes develop within the spore wall. Pollen grains (microgametophytes) mature from microspores, and ultimately produce sperm cells. Megagametophytes develop from megaspores and are retained within the ovule. Gymnosperms produce multiple archegonia, which produce the female gametes.[36]

During pollination, pollen grains are physically transferred between plants from the pollen cone to the ovule. Pollen is usually moved by wind or insects. Whole grains enter each ovule through a microscopic gap in the ovule coat (integument) called the micropyle. The pollen grains mature further inside the ovule and produce sperm cells. Two main modes of fertilization are found in gymnosperms. Cycads and Ginkgo have flagellated motile sperm[38] that swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm with no flagella that are moved along a pollen tube to the egg. After syngamy (joining of the sperm and egg cell), the zygote develops into an embryo (young sporophyte). More than one embryo is usually initiated in each gymnosperm seed. The mature seed comprises the embryo and the remains of the female gametophyte, which serves as a food supply, and the seed coat.[39]

Gymnosperms ordinarily reproduce by sexual reproduction, and only rarely express parthenogenesis.[40] Sexual reproduction in gymnosperms appears to be required for maintaining long-term genomic integrity.[40] Meiosis in sexual land plants provides a direct mechanism for repairing DNA in reproductive tissues.[40] The likely primary benefit of cross-pollination in gymnosperms, as in other eukaryotes, is that it allows the avoidance of inbreeding depression caused by the presence of recessive deleterious mutations.[41]

Genetics

The first published sequenced genome for any gymnosperm was the genome of Picea abies in 2013.[42]

Uses

Gymnosperms have major economic uses. Some, such as pine, fir, spruce, and cedar, are used for lumber, paper production, and resin. Some other common uses for gymnosperms are soap, varnish, nail polish, food, gum, and perfumes.[43]

References

  1. ↑ 1.0 1.1 1.2 (in en) The Ultimate Visual Family Dictionary. New Delhi: DK Pub.. 2012. pp. 122–125. ISBN 978-0-1434-1954-9. 
  2. ↑ Pandey, Saurabh; Moradi, Amir Bahram; Dovzhenko, Oleksandr; Touraev, Alisher; Palme, Klaus; Welsch, Ralf (2022-01-13). "Molecular Control of Sporophyte-Gametophyte Ontogeny and Transition in Plants". Frontiers in Plant Science 12. doi:10.3389/fpls.2021.789789. ISSN 1664-462X. PMID 35095963. Bibcode: 2022FrPS...1289789P. 
  3. ↑ Coniferae, Gnetophyta. "1 Relationships of Angiosperms to Other Seed Plants."
  4. ↑ Yang, Yong; Ferguson, David Kay; Liu, Bing; Mao, Kang-Shan; Gao, Lian-Ming; Zhang, Shou-Zhou; Wan, Tao; Rushforth, Keith et al. (2022-07-01). "Recent advances on phylogenomics of gymnosperms and a new classification" (in en). Plant Diversity 44 (4): 340–350. doi:10.1016/j.pld.2022.05.003. ISSN 2468-2659. PMID 35967253. Bibcode: 2022PlDiv..44..340Y. 
  5. ↑ 5.0 5.1 Raven, P.H. (2013). Biology of Plants. New York: W.H. Freeman and Co.. 
  6. ↑ Walas, Łukasz; Mandryk, Wojciech; Thomas, Peter A.; Tyrała-Wierucka, Żanna; Iszkuło, Grzegorz (2018-09-01). "Sexual systems in gymnosperms: A review" (in en). Basic and Applied Ecology 31: 1–9. doi:10.1016/j.baae.2018.05.009. ISSN 1439-1791. Bibcode: 2018BApEc..31....1W. https://www.sciencedirect.com/science/article/abs/pii/S1439179117304498. 
  7. ↑ "Sexual systems in gymnosperms: A review". Basic and Applied Ecology 31: 1–9. 2018. doi:10.1016/j.baae.2018.05.009. Bibcode: 2018BApEc..31....1W. http://eprints.keele.ac.uk/4961/1/29052018_1-s2.0-S1439179117304498-main.pdf. 
  8. ↑ Gehring, Catherine A.; Theimer, Tad C.; Whitham, Thomas G.; Keim, Paul (1998). "Ectomycorrhizal Fungal Community Structure of Pinyon Pines Growing in Two Environmental Extremes" (in en). Ecology 79 (5): 1562–1572. doi:10.1890/0012-9658(1998)079[1562:EFCSOP2.0.CO;2]. ISSN 1939-9170. https://esajournals.onlinelibrary.wiley.com/doi/abs/10.1890/0012-9658%281998%29079%5B1562%3AEFCSOP%5D2.0.CO%3B2. 
  9. ↑ Chang, Aimee Caye G.; Chen, Tao; Li, Nan; Duan, Jun (2019-08-14). "Perspectives on Endosymbiosis in Coralloid Roots: Association of Cycads and Cyanobacteria" (in English). Frontiers in Microbiology 10. doi:10.3389/fmicb.2019.01888. ISSN 1664-302X. PMID 31474965. 
  10. ↑ 10.0 10.1 "Gymnosperms on The Plant List". Theplantlist.org. http://www.theplantlist.org/browse/G/. 
  11. ↑ Li, Hong-Tao; Yi, Ting-Shuang; Gao, Lian-Ming; Ma, Peng-Fei; Zhang, Ting; Yang, Jun-Bo; Gitzendanner, Matthew A.; Fritsch, Peter W. et al. (May 2019). "Origin of angiosperms and the puzzle of the Jurassic gap" (in en). Nature Plants 5 (5): 461–470. doi:10.1038/s41477-019-0421-0. PMID 31061536. Bibcode: 2019NatPl...5..461L. 
  12. ↑ Morris, Jennifer L.; Puttick, Mark N.; Clark, James W.; Edwards, Dianne; Kenrick, Paul; Pressel, Silvia; Wellman, Charles H.; Yang, Ziheng et al. (2018-03-06). "The timescale of early land plant evolution". Proceedings of the National Academy of Sciences of the United States of America 115 (10): E2274–E2283. doi:10.1073/pnas.1719588115. PMID 29463716. Bibcode: 2018PNAS..115E2274M. 
  13. ↑ Jiao, Yuannian; Wickett, Norman J.; Ayyampalayam, Saravanaraj; Chanderbali, André S.; Landherr, Lena; Ralph, Paula E.; Tomsho, Lynn P.; Hu, Yi et al. (2011-04-10). "Ancestral polyploidy in seed plants and angiosperms". Nature 473 (7345): 97–100. doi:10.1038/nature09916. PMID 21478875. Bibcode: 2011Natur.473...97J. 
  14. ↑ Ollerton, J.; Coulthard, E. (2009). "Evolution of Animal Pollination". Science 326 (5954): 808–809. doi:10.1126/science.1181154. PMID 19892970. Bibcode: 2009Sci...326..808O. 
  15. ↑ Ren, D.; Labandeira, C.C.; Santiago-Blay, J.A.; Rasnitsyn, A. et al. (2009). "A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies". Science 326 (5954): 840–847. doi:10.1126/science.1178338. PMID 19892981. Bibcode: 2009Sci...326..840R. 
  16. ↑ Labandeira, Conrad C.; Yang, Qiang; Santiago-Blay, Jorge A.; Hotton, Carol L.; Monteiro, Antónia; Wang, Yong-Jie; Goreva, Yulia; Shih, ChungKun et al. (2016). "The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies". Proceedings of the Royal Society B: Biological Sciences 283 (1824). doi:10.1098/rspb.2015.2893. PMID 26842570. 
  17. ↑ Bond, W. J. (March 1989). "The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence". Biological Journal of the Linnean Society 36 (3): 227–249. doi:10.1111/j.1095-8312.1989.tb00492.x. 
  18. ↑ Fisher, Jack B.; Lindström, Anders; Marler, Thomas E. (2009-06-01). "Tissue Responses and Solution Movement After Stem Wounding in Six Cycas Species". HortScience 44 (3): 848–851. doi:10.21273/HORTSCI.44.3.848. ISSN 0018-5345. https://journals.ashs.org/hortsci/view/journals/hortsci/44/3/article-p848.xml. 
  19. ↑ Bell, Peter R.; Bell, Peter R.; Hemsley, Alan R. (2000). Green Plants: Their Origin and Diversity. Cambridge University Press. p. 247. ISBN 978-0-521-64673-4. https://books.google.com/books?id=HYkTvGq_RccC&q=armor%2520sclerenchymatous%2520leaf%2520bases&pg=PA247. 
  20. ↑ Cleal, Christopher J.; Thomas, Barry A. (2019). Introduction to Plant Fossils. Cambridge University Press. p. 179. ISBN 978-1-108-48344-5. https://books.google.com/books?id=KDGdDwAAQBAJ&q=%2522exclusively%2520underground%2C%2520like%2520a%2520tuber%2520%28e.g.%2520some%2520Zamia%2520species%29%2522&pg=PA179. 
  21. ↑ Mill, R. R. (2016-06-22). "A Monographic Revision of Retrophyllum (Podocarpaceae)". Edinburgh Journal of Botany 73 (2): 171–261. doi:10.1017/S0960428616000081. 
  22. ↑ A. Farjon, ed (2006). "Conifer database". http://www.catalogueoflife.org/show_database_details.php?database_name=Conifer+Database. 
  23. ↑ Campbell, Reece (2005). "Phylum Coniferophyta". Biology (7th ed.). p. 595. 
  24. ↑ Adams, Charles R.; Bamford, Katherine M.; Early, Mike P. (2012). Principles of Horticulture (6th ed.). Routledge. doi:10.4324/9780080969589. ISBN 978-0080969572. 
  25. ↑ "Conifer Life Cycle". https://www.treeguideuk.co.uk/conifer-life-cycle/. 
  26. ↑ Owens, J. N. (1998). "Pollination in conifers". Trends in Ecology & Evolution 13 (12): …. doi:10.1016/S1360-1385(98)01337-5. Bibcode: 1998TPS.....3..479O. https://www.sciencedirect.com/science/article/pii/S1360138598013375. Retrieved 25 January 2026. 
  27. ↑ Gilbert, Natasha (2010-09-28). "Threats to the world's plants assessed". Nature. doi:10.1038/news.2010.499. https://www.nature.com/articles/news.2010.499. 
  28. ↑ Leslie, Andrew B.Expression error: Unrecognized word "et". (2018). "An overview of extant conifer evolution from the perspective of the fossil record". American Journal of Botany 105 (9): 1531–1544. doi:10.1002/ajb2.1143. PMID 30157290. 
  29. ↑ Leslie, Andrew B. (2018). "ajb21143-sup-0004-AppendixS4". American Journal of Botany 105 (9): 1531–1544. doi:10.1002/ajb2.1143. PMID 30157290. https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf. 
  30. ↑ Stull, Gregory W.Expression error: Unrecognized word "et". (2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. PMID 34282286. Bibcode: 2021NatPl...7.1015S. https://www.nature.com/articles/s41477-021-00964-4. 
  31. ↑ Stull, Gregory W. (2021). main.dated.supermatrix.tree.T9.tre (Report). Figshare. doi:10.6084/m9.figshare.14547354.v1. https://figshare.com/articles/dataset/Gene_duplications_and_genomic_conflict_underlie_major_pulses_of_phenotypic_evolution_in_gymnosperms/14547354. 
  32. ↑ Cantino 2007.
  33. ↑ 33.0 33.1 Christenhusz, M.J.M.; Reveal, J.L.; Farjon, A.; Gardner, M.F.; Mill, R.R.; Chase, M.W. (2011). "A new classification and linear sequence of extant gymnosperms". Phytotaxa 19 (1): 55–70. doi:10.11646/phytotaxa.19.1.3. Bibcode: 2011Phytx..19...55C. http://www.mapress.com/phytotaxa/content/2011/f/pt00019p070.pdf. 
  34. ↑ Hilton, Jason; Bateman, Richard M. (January 2006). "Pteridosperms are the backbone of seed-plant phylogeny 1" (in en). The Journal of the Torrey Botanical Society 133 (1): 119–168. doi:10.3159/1095-5674(2006)133[119:PATBOS2.0.CO;2]. 
  35. ↑ Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1. Bibcode: 2016Phytx.261..201C. http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598. 
  36. ↑ 36.0 36.1 Samantha, Fowler; Rebecca, Roush; James, Wise (2013). "14.3 Seed Plants: Gymnosperms". Concepts of Biology. Houston, Texas: OpenStax. https://openstax.org/books/concepts-biology/pages/14-3-seed-plants-gymnosperms. Retrieved 31 March 2023. 
  37. ↑ Liu, Yang; Wang, Sibo; Li, Linzhou; Yang, Ting; Dong, Shanshan; Wei, Tong; Wu, Shengdan; Liu, Yongbo et al. (April 2022). "The Cycas genome and the early evolution of seed plants" (in en). Nature Plants 8 (4): 389–401. doi:10.1038/s41477-022-01129-7. ISSN 2055-0278. PMID 35437001. Bibcode: 2022NatPl...8..389L. 
  38. ↑ Southworth, Darlene; Cresti, Mauro (September 1997). "Comparison of flagellated and nonflagellated sperm in plants". American Journal of Botany 84 (9): 1301–1311. doi:10.2307/2446056. PMID 21708687. 
  39. ↑ Walters, Dirk R Walters Bonnie By (1996). Vascular plant taxonomy. Dubuque, Iowa: Kendall/Hunt Pub. Co.. p. 124. ISBN 978-0-7872-2108-9. https://archive.org/details/vascularplanttax00walt. 
  40. ↑ 40.0 40.1 40.2 Hörandl E. Apomixis and the paradox of sex in plants. Ann Bot. 2024 Mar 18:mcae044. doi: 10.1093/aob/mcae044. Epub ahead of print. PMID 38497809
  41. ↑ Charlesworth D, Willis JH. The genetics of inbreeding depression. Nat Rev Genet. 2009 Nov;10(11):783-96. doi: 10.1038/nrg2664. PMID 19834483
  42. ↑ Nystedt, B et al. (May 2013). "The Norway spruce genome sequence and conifer genome evolution". Nature 497 (7451): 579–584. doi:10.1038/nature12211. PMID 23698360. Bibcode: 2013Natur.497..579N. 
  43. ↑ Biswas, C.; Johri, B.M. (1997). "Economic Importance". The Gymnosperms. Springer, Berlin, Heidelberg. pp. 440–456. doi:10.1007/978-3-662-13164-0_23. ISBN 978-3-662-13166-4. https://link.springer.com/content/pdf/10.1007%2F978-3-662-13164-0.pdf. 

General bibliography

  • Cantino, Philip D.; Doyle, James A.; Graham, Sean W.; Judd, Walter S.; Olmstead, Richard G.; Soltis, Douglas E.; Soltis, Pamela S.; Donoghue, Michael J. (August 2007). "Towards a phylogenetic nomenclature of Tracheophyta". Taxon 56 (3): 822–846. doi:10.2307/25065864. Bibcode: 2007Taxon..56..822C. 
  • Gymnosperm Database
  • Gymnosperms on the Tree of Life
  • Christopher Nigel Page (2023).The Evolution of Arborescent Gymnosperms. Cambridge University Press. ISBN 978-1-009-26309-2. 2 volumes (volume one on the conifers of the Northern hemisphere and volume 2 on the conifers of the southern hemisphere).

Template:Acrogymnospermae classification Wikidata ☰ Q133712 entry



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