| Literature DB >> 36153445 |
Stem cambial variants of Taiwan lianas.
Sheng-Zehn Yang1, Po-Hao Chen2, Jian-Jhong Chen3.
Abstract
BACKGROUND: Cambium in lianas, responsible for secondary growth, develop diverse and diagnostic traits during the climbing phase. Studies on the cross-section of Taiwanese liana cambial variants are scarce. We collected multiple stem cross-sections from 287 liana species belonging to 52 families. Each sample was examined on five occasions, and the observations were documented.RESULTS: The results showed that approximately 22 cambial variants types were displayed in Taiwan lianas. Among these, axial vascular elements in radial segments were the most common, followed by the variants with the irregular conformation and intraxylary phloem. Based on our assessment, we provide the following identification features of a few families: Apocynaceae had intraxylary phloem; Convolvulaceae had intraxylary phloem combined with successive cambia; Lardizabalaceae, Menispermaceae, and Ranunculaceae possessed axial vascular elements in segments; Piperaceae had external primary vascular bundle cylinder combined with axial vascular elements in segments; Vitaceae had axial vascular elements in segments combined with irregular conformation. Axial vascular elements in segments and intraxylary phloem appeared in six or five combination types, showing that these two types combined with many variants are helpful for the identification of lianas. Two species, Momordica charantia var. abbreviata, and Momordica cochinchinensis had a cambium element in the outer cylinder of cortical bicollateral vascular bundles and formed directional layers of successive cambia.
CONCLUSIONS: Our study documented regular secondary growth with a single cambium in 36 species and cambial variants present in 16 species of Taiwanese lianas. Furthermore, we provide crucial baseline data on liana cambial variations, thereby improving our understanding of their morphology and identification.
© 2022. The Author(s).
Entities:
Keywords: Centrifugal xylem; Lianas; Parenchymatization; Secondary growth; Taiwan
Year: 2022 PMID： 36153445 PMCID： PMC9509506 DOI： 10.1186/s40529-022-00358-5
Source DB: PubMed Journal: Bot Stud ISSN： 1817-406X Impact factor: 2.673
Background
Materials and methods
Table 1
Cambial variants of each species in Taiwan lianas
| Families | Scientic name | Cambial variants |
|---|---|---|
| Acanthaceae | FD | |
| TE | ||
| TE | ||
| Actinidiaceae | RC | |
| RC | ||
| RC | ||
| RC | ||
| a | RC | |
| Amaranthaceae | SC | |
| Anacardiaceae | RC | |
| Annonaceae | RC | |
| RC | ||
| RC | ||
| Apocynaceae | TR | |
| a | TR | |
| TR | ||
| a
| TR | |
| TR | ||
| TR | ||
| TR, FC | ||
| a | TR, FC | |
| TR | ||
| TR | ||
| a
| TR | |
| TR | ||
| a
| TR | |
| TR, ES | ||
| TR, FC | ||
| a | TR | |
| TR | ||
| TR | ||
| Araliaceae | IC | |
| RC | ||
| a | RC | |
| Areacaceae | a | RC |
| a | RC | |
| Aristolochiaceae | a | VS |
| VS | ||
| VS | ||
| VS | ||
| Asteraceae | VS | |
| IC | ||
| FD | ||
| RC | ||
| VS | ||
| Basellaceae | VS, SC | |
| Bignoniaceae | FD | |
| FD | ||
| Cannabaceae | IC | |
| Capparaceae | a | RC |
| RC | ||
| Caprifoliaceae | RC | |
| RC | ||
| RC | ||
| Cecropiaceae | RC | |
| Celastraceae | FC | |
| VS | ||
| RC | ||
| FC | ||
| a | IC | |
| RC | ||
| Combretaceae | TR, FC | |
| Connaraceae | SC | |
| Convolvulaceae | a | TR, SC |
| a | TR, SC | |
| TR, SC | ||
| TR, SC | ||
| TR | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR | ||
| TR, SC | ||
| TR | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| TR, SC | ||
| Cucurbitaceae | IC, VS | |
| IC, VS | ||
| IC, VS | ||
| IC, VS, SC | ||
| IC, VS, SC | ||
| FD, VS, SC | ||
| FD, VS | ||
| Elaeagnaceae | RC | |
| RC | ||
| a | RC | |
| a | RC | |
| RC | ||
| Euphorbiaceae | IC | |
| Fabaceae | IC | |
| DX, IC | ||
| RC | ||
| RC | ||
| RC | ||
| RC | ||
| RC | ||
| IC | ||
| RC | ||
| RC | ||
| RC | ||
| FC | ||
| RC | ||
| RC | ||
| a | RC | |
| RC | ||
| a | FC | |
| RC | ||
| a | RC | |
| SC | ||
| SC | ||
| SC | ||
| ES, IC | ||
| RC | ||
| RC | ||
| RC | ||
| FC | ||
| RC | ||
| SC | ||
| RC | ||
| IC | ||
| RC | ||
| RC | ||
| RC | ||
| RC | ||
| SC | ||
| SC | ||
| IC | ||
| RC | ||
| Gesneriaceae | FC | |
| Heliotropiaceae | RC | |
| Hernandiaceae | FC | |
| Hydrangeaceae | RC | |
| a | RC | |
| RC | ||
| RC | ||
| Lardizabalaceae | VS | |
| VS | ||
| VS | ||
| VS | ||
| a
| VS | |
| Loganiaceae | FD, TR | |
| FD, TR | ||
| TE, TR | ||
| Malpighiaceae | RC | |
| FC | ||
| Melastomataceae | a | RC |
| a | TR | |
| Menispermaceae | VS | |
| VS, SC | ||
| a | VS | |
| VS | ||
| a | VS | |
| a | VS | |
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| Moraceae | IC | |
| IC | ||
| IC | ||
| IC | ||
| a | IC | |
| RC | ||
| RC | ||
| RC | ||
| Nyctaginaceae | SC | |
| TE | ||
| Oleaceae | IC | |
| RC | ||
| RC | ||
| RC | ||
| Opiliaceae | RC | |
| Pandanaceae | RC | |
| Passifloraceae | FC | |
| FC | ||
| RC | ||
| IC | ||
| FC | ||
| FC | ||
| Phyllanthaceae | FC | |
| Piperaceae | EP, VS | |
| EP, VS | ||
| EP, VS | ||
| EP, VS | ||
| a
| EP, VS | |
| a | EP, VS | |
| a | EP, VS | |
| a | EP, VS | |
| Polygonaceae | SC | |
| VS | ||
| FC | ||
| VS | ||
| Primulaceae | a | VS |
| a | VS | |
| VS | ||
| Ranunculaceae | VS | |
| VS | ||
| a | VS | |
| a | VS, FD | |
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| VS | ||
| a | VS | |
| VS | ||
| a | VS | |
| VS | ||
| VS | ||
| VS | ||
| Rhamnaceae | a | RC |
| a | RC | |
| RC | ||
| RC | ||
| a | RC | |
| a | RC | |
| RC | ||
| a | RC | |
| RC | ||
| Rosaceae | RC | |
| RC | ||
| RC | ||
| RC | ||
| IC | ||
| Rubiaceae | FC, IC | |
| DX | ||
| FC | ||
| FC | ||
| RC | ||
| RC | ||
| RC | ||
| a | RC | |
| IC, VS | ||
| IC, VS | ||
| RC | ||
| RC | ||
| RC | ||
| IC | ||
| RC | ||
| Rutaceae | RC | |
| RC | ||
| RC | ||
| Sabiaceae | VS | |
| a | VS | |
| Schisandraceae | RC | |
| RC | ||
| a | RC | |
| Solanaceae | TR | |
| TR | ||
| Vitaceae | IC | |
| IC | ||
| IC,VS, SC | ||
| IC,VS | ||
| VS | ||
| IC,VS | ||
| a | IC,VS | |
| IC,VS | ||
| IC,VS | ||
| IC,VS | ||
| VS | ||
| VS | ||
| IC,VS | ||
| a | IC,VS, SC | |
| a | IC,VS, SC | |
| a | IC,VS, SC | |
| IC,VS | ||
| IC | ||
| IC | ||
| a | IC | |
| IC | ||
| IC | ||
| 52 families | 287 species |
RC: stem cambium normal in production and round in conformation; VS: axial vascular elements in segments; IC: stem cambium normal in production but stem with irregular conformation; TR: intraxylary phloem; FC: furrowed xylem of cambium continuity; SC: successive cambia; FD: furrowed xylem of cambium discontinuity; TE: interxylary phloem; EP: external primary vascular cylinders; ES: external secondary vascular cylinders; DX: fissured stem/dispersed xylem
* Endemic species
Results and discussion
Patterns of cambial variants of Taiwan lianas
Table 2
Cambial variant types of lianas of different families in Taiwan
| Family | RC | VS | IC | TR | FC | SC | FD | TE | DX | TR + SC | IC + VS | EP + VS | FC + TR | FD + VS |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Acanthaceae | – | – | – | – | 1 | – | – | 2 | – | – | – | – | – | – |
| Actinidiaceae | 5 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Amaranthaceae | – | – | – | – | – | 1 | – | – | – | – | – | – | – | – |
| Anacardiaceae | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Annonaceae | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Apocynaceae | – | – | – | 14 | – | – | – | – | – | – | – | – | 3 | – |
| Araliaceae | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Areacaceae | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Aristolochiaceae | – | 4 | – | – | – | – | – | – | – | – | – | – | – | – |
| Asteraceae | 1 | 2 | 1 | – | – | – | 1 | – | – | – | – | – | – | – |
| Basellaceae | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Bignoniaceae | – | – | – | – | – | – | 2 | – | – | – | – | – | – | – |
| Cannabaceae | – | – | 1 | – | – | – | – | – | – | – | – | – | – | – |
| Capparaceae | 2 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Caprifoliaceae | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Cecropiaceae | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Celastraceae | 2 | 1 | 1 | – | 2 | – | – | – | – | – | – | – | – | – |
| Combretaceae | – | – | – | – | – | – | – | – | – | – | – | – | 1 | – |
| Connaraceae | – | – | – | – | – | 1 | – | – | – | – | – | – | – | – |
| Convolvulaceae | – | – | – | 4 | – | – | – | – | – | 20 | – | – | – | – |
| Cucurbitaceae | – | – | – | – | – | – | – | – | – | – | 3 | – | – | 1 |
| Elaeagnaceae | 5 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Euphorbiaceae | – | – | 1 | – | – | – | – | – | – | – | – | – | – | – |
| Fabaceae | 24 | – | 4 | – | 3 | 6 | – | – | – | – | – | – | – | – |
| Gesneriaceae | – | – | – | – | 1 | – | – | – | – | – | – | – | – | – |
| Heliotropiaceae | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Hernandiaceae | – | – | – | – | 1 | – | – | – | – | – | – | – | – | – |
| Hydrangeaceae | 4 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Lardizabalaceae | – | 5 | – | – | – | – | – | – | – | – | – | – | – | – |
| Loganiaceae | – | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Malpighiaceae | 1 | – | – | – | 1 | – | – | – | – | – | – | – | – | – |
| Melastomataceae | 1 | – | – | 1 | – | – | – | – | – | – | – | – | – | – |
| Menispermaceae | – | 13 | – | – | – | – | – | – | – | – | – | – | – | – |
| Moraceae | 4 | – | 4 | – | – | – | – | – | – | – | – | – | – | – |
| Nyctaginaceae | – | – | – | – | – | 1 | – | 1 | – | – | – | – | – | – |
| Oleaceae | 3 | – | 1 | – | – | – | – | – | – | – | – | – | – | – |
| Opiliaceae | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Pandanaceae | 1 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Passifloraceae | 1 | – | 1 | – | 4 | – | – | – | – | – | – | – | – | – |
| Phyllanthaceae | – | – | – | – | 1 | – | – | – | – | – | – | – | – | – |
| Piperaceae | – | – | – | – | – | – | – | – | – | – | – | 8 | – | – |
| Polygonaceae | – | 2 | – | – | 1 | 1 | – | – | – | – | – | – | – | – |
| Primulaceae | – | 3 | – | – | – | – | – | – | – | – | – | – | – | – |
| Ranunculaceae | – | 15 | – | – | – | – | – | – | – | – | – | – | – | 1 |
| Rhamnaceae | 9 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Rosaceae | 4 | – | 1 | – | – | – | – | – | – | – | – | – | – | – |
| Rubiaceae | 8 | – | 1 | – | 2 | – | – | – | 1 | – | 2 | – | – | – |
| Rutaceae | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Sabiaceae | – | 2 | – | – | – | – | – | – | – | – | – | – | – | – |
| Schisandraceae | 3 | – | – | – | – | – | – | – | – | – | – | – | – | – |
| Solanaceae | – | – | – | 2 | – | – | – | – | – | – | – | – | – | – |
| Vitaceae | – | 3 | 7 | – | – | – | – | – | – | – | 8 | – | – | – |
| Sum of family | 25 | 11 | 11 | 4 | 10 | 5 | 2 | 2 | 1 | 1 | 3 | 1 | 2 | 2 |
| Sum of species | 96 | 50 | 23 | 21 | 17 | 10 | 3 | 3 | 1 | 20 | 13 | 8 | 4 | 2 |
| Percentage % | 33.4 | 17.4 | 8.0 | 7.3 | 5.5 | 3.5 | 1.0 | 1.0 | 0.3 | 7.0 | 4.5 | 2.8 | 1.4 | 0.7 |
RC, stem cambium normal in production and round in conformation; VS, axial vascular elements in segments; IC, stem cambium normal in production but stem with irregular conformation; TR, intraxylary phloem; FC, furrowed xylem of cambium continuity; SC, successive cambia; FD, furrowed xylem of cambium discontinuity; TE, interxylary phloem; EP, external primary vascular cylinders; ES, external secondary vascular cylinders; DX, fissured stem/dispersed xylem; SP No., species number; Types No., cambium variant types in each family, RC not included
Fig. 1Single cambial variant of stem cross section. A Axial vascular elements in segments: Pericampylus formosanus. B Irregular in conformation: Jasminum lanceolarium. C Intraxylary phloem: Melodinus angustifolius. D Furrowed xylem of cambium continuity: Morinda parvifolia. E Successive cambia: Mucuna macrocarpa. F Furrowed xylem of cambium discontinuity: Pyrostegia venusta. RD, ray dilatation, PI, pith, XY, xylem, PH, phloem, IC, irregular conformation, TR, intraxylary phloem, CO, cork, FC, furrowed xylem of cambium continuity, PA, parenchyma, 1st XY, first xylem, 2nd XP, second xylem, 3rd XP, third xylem, 4th XY, forth xylem, 1st PH, first phloem, 2nd PH, second phloem, 3rd PH, third phloem, 4th PH, forth phloem, FD, furrowed xylem of cambium discontinuity, PHW, phloem wedges-like
Fig. 2Single cambial variant or combination of two cambial variants of stem cross section. A Interxylary phloem (enlargement): Thunbergia grandiflora. B Intraxylary phloem (enlargement) combined with successive cambia: Distimake tuberosus. C Irregular conformation combined with axial vascular elements in segments: Tetrastigma obtectum var. glabrum. D External primary vascular cylinders combined with axial vascular elements in segments: Piper kadsura. E Furrowed xylem of cambium continuity combined with intraxylary phloem: Gymnema sylvestre. F Furrowed xylem of cambium discontinuity combined with axial vascular elements in segments: Clematis gouriana subsp. lishanensis. PI, pith, TE, interxylary phloem, TR, intraxylary phloem, 1st XY, first xylem, 2nd XP, second xylem, 1st PH, first phloem, RA, ray, XY, xylem, PH, phloem, SCL, sclenchyma, MU, mucilage canal, ME, medullary vascular bundles, PE, peripheral vascular bundles, FD, furrowed xylem of cambium discontinuity, FC, furrowed xylem of cambium continuity, PW, phloem wedges
Fig. 4Combination of three cambial variants or single cambial variant of stem cross section. A Successive cambia combined with axial vascular elements in segments and irregular conformation: Tetrastigma formosanum. B Axial vascular elements in segments combined with successive cambia: Anredera cordifolia. C External secondary vascular cylinders (empty star) combined with axial vascular elements in segments and successive cambia: Momordica charantia var. abbreviata. D Fissured stem/dispersed xylem combined: Dinetia hedyotidea. E Furrowed xylem of cambium continuity: Passiflora edulis. F Irregular in conformation combined with axial vascular elements in segments and successive cambia: Cayratia corniculata. PI, pith, PIC, pith cavity, RA, ray, PW, phloem wedges, 1st XY, first xylem, 2nd XY, second xylem, 3rd XY, third xylem, 1st PH, first phloem, 2nd PH, second phloem, 3rd PH, third phloem, 2nd VA, second vascular bundle, IC, irregular conformation, PA, parenchyma proliferation, DX, fissured stem/dispersed xylem
Fig. 3Combination of two cambial variants of stem cross section. A Furrowed xylem of cambium discontinuity combined with intraxylary phloem: Gardneria multiflora. B Furrowed xylem of cambium continuity combined with irregular conformation; Coptosapelta diffusa. C External secondary vascular cylinders combined with intraxylary phloem: Trachelospermum gracilipes. D Interxylary phloem combined with intraxylary phloem: Strychnos cathayensis. E External secondary vascular cylinders combined with irregular conformation: Lablab purpureus. F Fissured stem/dispersed xylem combined with irregular conformation: Bauhinia championii. PI, pith, FC, furrowed xylem of cambium continuity, PW, phloem wedges, IC, irregular conformation, PA, parenchyma, DX, fissured stem/ dispersed xylem, TR, intraxylary phloem, ES, external secondary vascular cylinders, XY, xylem, PH, phloem, TE, interxylary phloem
Table 3
List of orders and families of Taiwan lianas that possess the cambial variants
| Orders | Families | Cambial variants (this study) | Angyalossy et al. ( |
|---|---|---|---|
| Asterales | Asteraceae | VS, FC, FD | VS, FX |
| Apiales | Araliaceae | RC | – |
| Arecales | Areacaceae | RC | – |
| Austrobaileyales | Schisandraceae | RC | – |
| Boraginales | Heliotropiaceae | RC | – |
| Brassicales | Capparaceae | RC | – |
| Caryophyllales | Amaranthaceae | SC | SC |
| Nyctaginaceae | TE, SC | SC | |
| Polygonaceae | SC, VS, FC | IC, SC, TR | |
| Basellaceae | VS, TR, SC | – | |
| Cornales | Hydrangeaceae | RC | – |
| Celastrales | Celastraceae | FC, VS, IC | FX, IC, SC,TE |
| Cucurbitales | Cucurbitaceae | IC + VS, VS + IS + SC, FD + VS, FD + VS + SC | SC, TE, VS |
| Dipsacales | Caprifoliaceae | RC | – |
| Ericales | Actinidiaceae | RC | – |
| Primulaceae | VS | – | |
| Fabales | Fabaceae | IC, DX + IC, FC, SC, ES | SC, TE, IC, FX, DX, TR |
| Gentianales | Apocynaceae | TR, TR + FC, TR + ES | IC, SC, TE, VS, FX |
| Loganiaceae | TR + FD, TR + TE | TE | |
| Rubiaceae | FC + IC, IC, DX + TR + IC, IC + VS | FX, IC | |
| Lamiales | Acanthaceae | FD, TE | SC, TE, DX |
| Bignoniaceae | FD | FX, DX | |
| Gesneriaceae | FC | – | |
| Oleaceae | IC | – | |
| Laurales | Hernandiaceae | FC | – |
| Magnoliales | Annonaceae | RC | – |
| Malpighiales | Euphorbiaceae | IC | ES |
| Malpighiaceae | FC | FX, DX, SC, TE, IC | |
| Passifloraceae | FC | SC, FX | |
| Phyllanthaceae | FC | – | |
| Myrtales | Combretaceae | TR + FC | TR, TE, ES, IC |
| Oxalidales | Connaraceae | SC | SC |
| Pandanales | Pandanaceae | RC | – |
| Piperales | Aristolochiaceae | VS | VS, IC |
| Piperaceae | EP + VS | VS, IC, EP | |
| Proteales | Sabiaceae | VS | – |
| Ranunculales | Menispermaceae | VS, VS + SC | SC, VS, IC, TR |
| Ranunculaceae | VS, VS + FD | VS | |
| Lardizabalaceae | VS | – | |
| Rosales | Cannabaceae | IC | – |
| Cecropiaceae | RC | – | |
| Elaeagnaceae | RC | – | |
| Rhamnaceae | RC | – | |
| Rosaceae | RC | – | |
| Moraceae | IC | IC | |
| Santalales | Opiliaceae | RC | – |
| Solanales | Convolvulaceae | TR, TR + SC | SC, TE, FX, DX, TR |
| Solanaceae | TR | – | |
| Vitales | Vitaceae | IC, VS, IC + VS + SC, VS + IC | VS, IC, SC, DX |
| 18 orders | 49 families |
RC, stem cambium normal in production and round in conformation; VS, axial vascular elements in segments; IC, stem cambium normal in production but stem with irregular conformation; TR, intraxylary phloem; FC, furrowed xylem of cambium continuity; SC, successive cambia; FD, furrowed xylem of cambium discontinuity; TE, interxylary phloem; EP, external primary vascular cylinders; ES, external secondary vascular cylinders; DX, fissured stem/dispersed xylem
Cambial variants of Taiwan lianas in each family or genus
Fig. 5The bicollateral vascular bundle (Cucurbitaceae) with outer and inner phloem at both ends. White stars are inner and outer phloem
Cambial variants of Taiwan lianas in each order
Conclusion
4 in total
1. The rise and evolution of the cambial variant in Bignonieae (Bignoniaceae).
Journal: Evol Dev Date: 2009 Sep-Oct Impact factor: 1.930
2. Moving with climbing plants from Charles Darwin's time into the 21st century.
Journal: Am J Bot Date: 2009-07 Impact factor: 3.844
3. Phloem wedges in Malpighiaceae: origin, structure, diversification, and systematic relevance.
Journal: Evodevo Date: 2022-04-28 Impact factor: 3.569
4. Cambial variations of Piper (Piperaceae) in Taiwan.
Journal: Bot Stud Date: 2017-03-29 Impact factor: 2.787
4 in total