Plant morphology

Short description: Study of the structure of plants

Inflorescences emerging from protective coverings of Angelica gigas

Plant morphology or phytomorphology is the study of the physical form and external structure of plants.[1] This is usually considered distinct from plant anatomy,[1] which is the study of the internal structure of plants, especially at the microscopic level.[2] Plant morphology is useful in the visual identification of plants. Recent studies in molecular biology started to investigate the molecular processes involved in determining the conservation and diversification of plant morphologies. In these studies, transcriptome conservation patterns were found to mark crucial ontogenetic transitions during the plant life cycle which may result in evolutionary constraints limiting diversification.[3]

## Contents

* 1 Scope
* 2 A comparative science
* 2.1 Homology
* 2.2 Convergence
* 3 Vegetative and reproductive characteristics
* 3.1 Use in identification
* 3.2 Alternation of generations
* 3.3 Pigmentation in plants
* 4 Morphology in development
* 4.1 Growth
* 4.2 Morphological variation
* 4.3 Evolution of plant morphology
* 4.3.1 Positional effects
* 4.3.2 Environmental effects
* 4.3.2.1 Temperature
* 4.3.3 Juvenility
* 5 Modern Innovations
* 6 See also
* 7 References
* 8 External links

## Scope

Asclepias syriaca showing complex morphology of the flowers. Looking up into the branch structure of a Pinus sylvestris tree

File:Travel Deep Inside a Leaf - Annotated Version - California Academy of Sciences.webm

Plant morphology "represents a study of the development, form, and structure of plants, and, by implication, an attempt to interpret these on the basis of similarity of plan and origin".[4] There are four major areas of investigation in plant morphology, and each overlaps with another field of the biological sciences.[1]

First of all, morphology is comparative, meaning that the morphologist examines structures in many different plants of the same or different species, then draws comparisons and formulates ideas about similarities. When structures in different species are believed to exist and develop as a result of common, inherited genetic pathways, those structures are termed homologous. For example, the leaves of pine, oak, and cabbage all look very different, but share certain basic structures and arrangement of parts. The homology of leaves is an easy conclusion to make. The plant morphologist goes further, and discovers that the spines of cactus also share the same basic structure and development as leaves in other plants, and therefore cactus spines are homologous to leaves as well. This aspect of plant morphology overlaps with the study of plant evolution and paleobotany.[5]

Secondly, plant morphology observes both the vegetative (somatic) structures of plants, as well as the reproductive structures. The vegetative structures of vascular plants includes the study of the shoot system, composed of stems and leaves, as well as the root system. The reproductive structures are more varied, and are usually specific to a particular group of plants, such as flowers and seeds, fern sori, and moss capsules. The detailed study of reproductive structures in plants led to the discovery of the alternation of generations found in all plants and most algae. This area of plant morphology overlaps with the study of biodiversity and plant systematics.[4]

Thirdly, plant morphology studies plant structure at a range of scales. At the smallest scales are ultrastructure, the general structural features of cells visible only with the aid of an electron microscope, and cytology, the study of cells using optical microscopy. At this scale, plant morphology overlaps with plant anatomy as a field of study. At the largest scale is the study of plant growth habit, the overall architecture of a plant. The pattern of branching in a tree will vary from species to species, as will the appearance of a plant as a tree, herb, or grass.[6]

Fourthly, plant morphology examines the pattern of development, the process by which structures originate and mature as a plant grows. While animals produce all the body parts they will ever have from early in their life, plants constantly produce new tissues and structures throughout their life. A living plant always has embryonic tissues. The way in which new structures mature as they are produced may be affected by the point in the plant's life when they begin to develop, as well as by the environment to which the structures are exposed. A morphologist studies this process, the causes, and its result. This area of plant morphology overlaps with plant physiology and ecology.

## A comparative science

A plant morphologist makes comparisons between structures in many different plants of the same or different species. Making such comparisons between similar structures in different plants tackles the question of why the structures are similar. It is quite likely that similar underlying causes of genetics, physiology, or response to the environment have led to this similarity in appearance. The result of scientific investigation into these causes can lead to one of two insights into the underlying biology:[7]

1. Homology \- the structure is similar between the two species because of shared ancestry and common genetics.
2. Convergence \- the structure is similar between the two species because of independent adaptation to common environmental pressures.

### Homology

### Convergence

Main article: Biology:Convergent evolution

## Vegetative and reproductive characteristics

A diagram representing a "typical" eudicot.

Plant morphology treats both the vegetative structures of plants, as well as the reproductive structures.

The vegetative (somatic) structures of vascular plants include two major organ systems: (1) a shoot system, composed of stems and leaves, and (2) a root system. These two systems are common to nearly all vascular plants, and provide a unifying theme for the study of plant morphology.

By contrast, the reproductive structures are varied, and are usually specific to a particular group of plants. Structures such as flowers and fruits are only found in the angiosperms; sori are only found in ferns; and seed cones are only found in conifers and other gymnosperms. Reproductive characters are therefore regarded as more useful for the classification of plants than vegetative characters.

### Use in identification

Main article: Biology:Identification key

Plant biologists use morphological characters of plants which can be compared, measured, counted and described to assess the differences or similarities in plant taxa and use these characters for plant identification, classification and descriptions.

When characters are used in descriptions or for identification they are called diagnostic or key characters which can be either qualitative and quantitative.

1. Quantitative characters are morphological features that can be counted or measured for example a plant species has flower petals 10–12 mm wide.
2. Qualitative characters are morphological features such as leaf shape, flower color or pubescence.

Both kinds of characters can be very useful for the identification of plants.

### Alternation of generations

Main article: Biology:Alternation of generations

### Pigmentation in plants

## Morphology in development

Main article: Biology:Plant development

The process of development in plants is fundamentally different from that seen in vertebrate animals. When an animal embryo begins to develop, it will very early produce all of the body parts that it will ever have in its life. When the animal is born (or hatches from its egg), it has all its body parts and from that point will only grow larger and more mature. By contrast, plants constantly produce new tissues and structures throughout their life from meristems[8] located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues.

### Growth

Once the embryo germinates from its seed or parent plant, it begins to produce additional organs (leaves, stems, and roots) through the process of organogenesis. New roots grow from root meristems located at the tip of the root, and new stems and leaves grow from shoot meristems located at the tip of the shoot.[9] Branching occurs when small clumps of cells left behind by the meristem, and which have not yet undergone cellular differentiation to form a specialised tissue, begin to grow form the tip of a new root or shoot. Growth from any such meristem at the tip of a root or shoot is termed primary growth and results in the lengthening of that root or shoot. Secondary growth results in widening of a root or shoot from divisions of cells in a cambium.[10]

In addition to growth by cell division, a plant may grow through cell elongation. This occurs when individual cells or groups of cells grow longer. Not all plant cells will grow to the same length. When cells on one side of a stem grow longer and faster than cells on the other side, the stem will bend to the side of the slower growing cells as a result. This directional growth can occur via a plant's response to a particular stimulus, such as light (phototropism), gravity (gravitropism), water, (hydrotropism), and physical contact (thigmotropism).[11]

Plant growth and development are mediated by specific plant hormones and plant growth regulators (PGRs) (Ross et al. 1983).[12] Endogenous hormone levels are influenced by plant age, cold hardiness, dormancy, and other metabolic conditions; photoperiod, drought, temperature, and other external environmental conditions; and exogenous sources of PGRs, e.g., externally applied and of rhizospheric origin.[13]

### Morphological variation

Plants exhibit natural variation in their form and structure. While all organisms vary from individual to individual, plants exhibit an additional type of variation. Within a single individual, parts are repeated which may differ in form and structure from other similar parts. This variation is most easily seen in the leaves of a plant, though other organs such as stems and flowers may show similar variation. There are three primary causes of this variation: positional effects, environmental effects, and juvenility.[14]

### Evolution of plant morphology

Transcription factors and transcriptional regulatory networks play key roles in plant morphogenesis and their evolution. During plant landing, many novel transcription factor families emerged and are preferentially wired into the networks of multicellular development, reproduction, and organ development, contributing to more complex morphogenesis of land plants.[15]

#### Positional effects

Variation in leaves from the giant ragweed illustrating positional effects. The lobed leaves come from the base of the plant, while the unlobed leaves come from the top of the plant.

Although plants produce numerous copies of the same organ during their lives, not all copies of a particular organ will be identical. There is variation among the parts of a mature plant resulting from the relative position where the organ is produced. For example, along a new branch the leaves may vary in a consistent pattern along the branch. The form of leaves produced near the base of the branch will differ from leaves produced at the tip of the plant, and this difference is consistent from branch to branch on a given plant and in a given species. This difference persists after the leaves at both ends of the branch have matured, and is not the result of some leaves being younger than others.[16][17]

#### Environmental effects

The way in which new structures mature as they are produced may be affected by the point in the plant's life when they begin to develop, as well as by the environment to which the structures are exposed. This can be seen in aquatic plants.[18]

##### Temperature

Temperature has a multiplicity of effects on plants depending on a variety of factors, including the size and condition of the plant and the temperature and duration of exposure. The smaller and more succulent the plant, the greater the susceptibility to damage or death from temperatures that are too high or too low. Temperature affects the rate of biochemical and physiological processes, rates generally (within limits) increasing with temperature. However, the Van't Hoff relationship for monomolecular reactions (which states that the velocity of a reaction is doubled or trebled by a temperature increase of 10 °C) does not strictly hold for biological processes, especially at low and high temperatures.[19]

When water freezes in plants, the consequences for the plant depend very much on whether the freezing occurs intracellularly (within cells) or outside cells in intercellular (extracellular) spaces.[20] Intracellular freezing usually kills the cell regardless of the hardiness of the plant and its tissues.[21] Intracellular freezing seldom occurs in nature, but moderate rates of decrease in temperature, e.g., 1 °C to 6 °C/hour, cause intercellular ice to form, and this "extraorgan ice"[22] may or may not be lethal, depending on the hardiness of the tissue.

At freezing temperatures, water in the intercellular spaces of plant tissues freezes first, though the water may remain unfrozen until temperatures fall below 7 °C.[20] After the initial formation of ice intercellularly, the cells shrink as water is lost to the segregated ice. The cells undergo freeze-drying, the dehydration being the basic cause of freezing injury.[23]

The rate of cooling has been shown to influence the frost resistance of tissues,[24] but the actual rate of freezing will depend not only on the cooling rate, but also on the degree of supercooling and the properties of the tissue.[25] Sakai (1979a)[24] demonstrated ice segregation in shoot primordia of Alaskan white and black spruces when cooled slowly to 30 °C to -40 °C. These freeze-dehydrated buds survived immersion in liquid nitrogen when slowly rewarmed. Floral primordia responded similarly. Extraorgan freezing in the primordia accounts for the ability of the hardiest of the boreal conifers to survive winters in regions when air temperatures often fall to -50 °C or lower.[22] The hardiness of the winter buds of such conifers is enhanced by the smallness of the buds, by the evolution of faster translocation of water, and an ability to tolerate intensive freeze dehydration. In boreal species of Picea and Pinus, the frost resistance of 1-year-old seedlings is on a par with mature plants,[26] given similar states of dormancy.

#### Juvenility

Juvenility in a seedling of European beech. There is a marked difference in shape between the first dark green "seed leaves" and the lighter second pair of leaves.

The organs and tissues produced by a young plant, such as a seedling, are often different from those that are produced by the same plant when it is older. This phenomenon is known as juvenility or heteroblasty. For example, young trees will produce longer, leaner branches that grow upwards more than the branches they will produce as a fully grown tree. In addition, leaves produced during early growth tend to be larger, thinner, and more irregular than leaves on the adult plant. Specimens of juvenile plants may look so completely different from adult plants of the same species that egg-laying insects do not recognise the plant as food for their young. Differences are seen in rootability and flowering and can be seen in the same mature tree. Juvenile cuttings taken from the base of a tree will form roots much more readily than cuttings originating from the mid to upper crown. Flowering close to the base of a tree is absent or less profuse than flowering in the higher branches especially when a young tree first reaches flowering age.[27]

The transition from early to late growth forms is referred to as 'vegetative phase change', but there is some disagreement about terminology.[28]

## Modern Innovations

Rolf Sattler has revised fundamental concepts of comparative morphology such as the concept of homology. He emphasised that homology should also include partial homology and quantitative homology.[29][30] This leads to a continuum morphology that demonstrates a continuum between the morphological categories of root, shoot, stem (caulome), leaf (phyllome), and hair (trichome). How intermediates between the categories are best described has been discussed by Bruce K. Kirchoff et al.[31] A recent study conducted by Salk Institute extracted coordinates corresponding to each plant's base and leaves in 3D space. When plants on the graph were placed according to their actual nutrient travel distances and total branch lengths, the plants fell almost perfectly on the Pareto curve. "This means the way plants grow their architectures also optimises a very common network design tradeoff. Based on the environment and the species, the plant is selecting different ways to make tradeoffs for those particular environmental conditions."[32]

Honoring Agnes Arber, author of the partial-shoot theory of the leaf, Rutishauser and Isler called the continuum approach Fuzzy Arberian Morphology (FAM). "Fuzzy" refers to fuzzy logic, "Arberian" to Agnes Arber. Rutishauser and Isler emphasised that this approach is not only supported by many morphological data but also by evidence from molecular genetics.[33] More recent evidence from molecular genetics provides further support for continuum morphology. James (2009) concluded that "it is now widely accepted that... radiality [characteristic of most stems] and dorsiventrality [characteristic of leaves] are but extremes of a continuous spectrum. In fact, it is simply the timing of the KNOX gene expression!."[34] Eckardt and Baum (2010) concluded that "it is now generally accepted that compound leaves express both leaf and shoot properties."[35]

Process morphology describes and analyses the dynamic continuum of plant form. According to this approach, structures do not have process(es), they are process(es).[36][37][38] Thus, the structure/process dichotomy is overcome by "an enlargement of our concept of 'structure' so as to include and recognise that in the living organism it is not merely a question of spatial structure with an 'activity' as something over or against it, but that the concrete organism is a spatio-temporal structure and that this spatio-temporal structure is the activity itself".[39]

For Jeune, Barabé and Lacroix, classical morphology (that is, mainstream morphology, based on a qualitative homology concept implying mutually exclusive categories) and continuum morphology are sub-classes of the more encompassing process morphology (dynamic morphology).[40]

Classical morphology, continuum morphology, and process morphology are highly relevant to plant evolution, especially the field of plant evolutionary biology (plant evo-devo) that tries to integrate plant morphology and plant molecular genetics.[41] In a detailed case study on unusual morphologies, Rutishauser (2016) illustrated and discussed various topics of plant evo-devo such as the fuzziness (continuity) of morphological concepts, the lack of a one-to-one correspondence between structural categories and gene expression, the notion of morphospace, the adaptive value of bauplan features versus patio ludens, physiological adaptations, hopeful monsters and saltational evolution, the significance and limits of developmental robustness, etc.[42] Rutishauser (2020) discussed the past and future of plant evo-devo.[43] Our conception of the gynoecium and the search for a fossil ancestor of Angiosperms changes fundamentally from the perspective of evo-devo.[44]

Whether we like it or not, morphological research is influenced by philosophical assumptions such as either/or logic, fuzzy logic, structure/process dualism or its transcendence. And empirical findings may influence the philosophical assumptions. Thus there are interactions between philosophy and empirical findings. These interactions are the subject of what has been referred to as philosophy of plant morphology.[45]

One important and unique event in plant morphology of the 21st century was the publication of Principles of Plant Morphology by Donald R. Kaplan, edited by Chelsea D. Specht (2022)[46] It is a well illustrated volume of 1305 pages in a very large format that presents a wealth of morphological data. Unfortunately, all of these data are only interpreted in terms of classical morphology and the qualitative homology concept, disregarding modern conceptional innovations.[47] Including continuum and process morphology as well as molecular genetics would provide an enlarged scope.[48]

An even more important event was the publication of a book by Regine Claβen-Bockhoff (2024) Die Pflanze: Morphologie, Entwicklung und Evolution von Vielfalt (The Plant: Morphology, Development and Evolution of Diversity).[49][50] Like Kaplan's book, this book is very comprehensive (over a thousand pages) and beautifully illustrated (she worked with two illustrators), but unlike Kaplan's book, her book presents major conceptual innovations. Although, for the vegetative region, she accepts the categories of classical morphology, contrary to Kaplan, she recognizes that not all structures can be pressed into these categories. For flowers, she abandoned the classical framework altogether. Instead of interpreting the flower as a modified short shoot (as posited by classical morphology), she proposed that flowers are sporangia bearing units so that stamens and carpels are sporangiophores, which are considered 'de novo' structures not necessarily homologous with vegetative leaves.[50]

Rolf Sattler proposed an Articulation Morphology.[51] It is based on the open growth of plants, which occurs through ramification that leads to articulation - the formation of articles between successive ramifications or after a single ramification. Thus, the plant is seen as an articulated whole, consisting of articles. The concept of the article encompasses the continuum between morphological categories and different levels of organization, from organ systems to organs, parts of organs and trichomes. Furthermore, it applies to all plants, from algae to bryophytes and vascular plants. In articulation morphology, the central and most basic concept is no longer morphological homology but transformation: transformation of ramification and articulation. This changes the most basic questions we ask. Instead of asking questions about morphological homology, we ask how ramification and articulation have changed during development and evolution. For this reason, articulation morphology may be considered a new paradigm of plant morphology. It changes fundamentally our way of thinking about morphology and morphological investigation.

## See also

* Glossary of plant morphology
* Plant anatomy
* Plant identification
* Plant physiology
* Plant evolutionary developmental biology
* Taxonomy

## References

1. ↑ 1.0 1.1 1.2 Raven, P. H., R. F. Evert, & S. E. Eichhorn. Biology of Plants, 7th ed., page 9. (New York: W. H. Freeman, 2005). ISBN 0-7167-1007-2.
2. ↑ Evert, Ray Franklin and Esau, Katherine (2006) Esau's Plant anatomy: meristems, cells, and tissues of the plant body - their structure, function and development Wiley, Hoboken, New Jersey, page xv, ISBN 0-471-73843-3
3. ↑ Drost, Hajk-Georg; Bellstaedt, Julia; Ó'Maoiléidigh, Diarmuid S.; Silva, Anderson T.; Gabel, Alexander; Weinholdt, Claus; Ryan, Patrick T.; Dekkers, Bas J.W. et al. (2016-02-23). "Post-embryonic Hourglass Patterns Mark Ontogenetic Transitions in Plant Development" (in en). Molecular Biology and Evolution 33 (5): 1158–1163. doi:10.1093/molbev/msw039. PMID 26912813. PMC 4839224. https://academic.oup.com/mbe/article/33/5/1158/2580081.
4. ↑ 4.0 4.1 Harold C. Bold, C. J. Alexopoulos, and T. Delevoryas. Morphology of Plants and Fungi, 5th ed., page 3. (New York: Harper-Collins, 1987). ISBN 0-06-040839-1.
5. ↑ Miravel-Gabriel, Cristina Betzabeth; Koeth, Ryan; Marsch-Martínez, Nayelli; Hernández-Hernández, Tania (2024-12-04). "Are Cactus Spines Modified Leaves? Morphological and Anatomical Characterization of Saguaro Seedlings (Carnegiea gigantea) with Special Focus on Aerial Organ Primordia" (in en). Plants 13 (23): 3406. doi:10.3390/plants13233406. ISSN 2223-7747. PMID 39683199. Bibcode: 2024Plnts..13.3406M.
6. ↑ Hallé, Francis; Oldeman, Roelof A. A.; Tomlinson, Philip B. (1978). "Tropical Trees and Forests". Springer. doi:10.1007/978-3-642-81190-6. ISBN 978-3-642-81192-0. https://link.springer.com/book/10.1007/978-3-642-81190-6?error=cookies_not_supported&code=7c6dbc3d-88ed-4b5f-af2d-402a26edfbee.
7. ↑ Sattler, Rolf (1992). "Process morphology: structural dynamics in development and evolution". Canadian Journal of Botany 70 (4): 708–714. doi:10.1139/b92-091. ISSN 0008-4026. Bibcode: 1992CaJB...70..708S.
8. ↑ Bäurle, I; Laux, T (2003). "Apical meristems: The plant's fountain of youth". BioEssays 25 (10): 961–70. doi:10.1002/bies.10341. PMID 14505363. Bibcode: 2003BiEss..25..961B. Review.
9. ↑ Brand, U; Hobe, M; Simon, R (2001). "Functional domains in plant shoot meristems". BioEssays 23 (2): 134–41. doi:10.1002/1521-1878(200102)23:2<134::AID-BIES1020>3.0.CO;2-3. PMID 11169586. Review.
10. ↑ Barlow, P (2005). "Patterned cell determination in a plant tissue: The secondary phloem of trees". BioEssays 27 (5): 533–41. doi:10.1002/bies.20214. PMID 15832381.
11. ↑ Esmon, C. Alex; Pedmale, Ullas V.; Liscum, Emmanuel (2005). "Plant tropisms: providing the power of movement to a sessile organism". The International Journal of Developmental Biology 49 (5–6): 665–674. doi:10.1387/ijdb.052028ce. ISSN 0214-6282. PMID 16096973.
12. ↑ Ross, S. D.; Pharis, R. P.; Binder, W. D. (2018-01-18), Nickell, Louis G., ed., "Growth Regulators and Conifers: Their Physiology and Potential Uses in Forestry" (in en), Plant Growth Regulating Chemicals (CRC Press): pp. 35–78, doi:10.1201/9781351075749-2, ISBN 978-1-351-07574-9, https://www.taylorfrancis.com/books/9781351084192/chapters/10.1201/9781351075749-2, retrieved 2026-05-05
13. ↑ Vanstraelen, Marleen; Benková, Eva (2012-11-10). "Hormonal Interactions in the Regulation of Plant Development" (in en). Annual Review of Cell and Developmental Biology 28 (2012): 463–487. doi:10.1146/annurev-cellbio-101011-155741. ISSN 1081-0706. PMID 22856461. https://www.annualreviews.org/content/journals/10.1146/annurev-cellbio-101011-155741.
14. ↑ Poethig, R. Scott (2013), "Vegetative Phase Change and Shoot Maturation in Plants" (in en), Current Topics in Developmental Biology (Elsevier) 105: 125–152, doi:10.1016/B978-0-12-396968-2.00005-1, ISBN 978-0-12-396968-2, PMID 23962841
15. ↑ Jin JP (July 2015). "An Arabidopsis transcriptional regulatory map reveals distinct functional and evolutionary features of novel transcription factors". Molecular Biology and Evolution 32 (7): 1767–1773. doi:10.1093/molbev/msv058. PMID 25750178.
16. ↑ Breuer, Christian; Braidwood, Luke; Sugimoto, Keiko (2014). "Endocycling in the path of plant development" (in en). Current Opinion in Plant Biology 17: 78–85. doi:10.1016/j.pbi.2013.11.007. ISSN 1369-5266. PMID 24507498. Bibcode: 2014COPB...17...78B. https://botanapp.com/.
17. ↑ Critchfield, William B. (1960). "Leaf Dimorphism in Populus Trichocarpa" (in en). American Journal of Botany 47 (8): 699–711. doi:10.1002/j.1537-2197.1960.tb07154.x. ISSN 0002-9122. Bibcode: 1960AmJB...47..699C. https://bsapubs.onlinelibrary.wiley.com/doi/10.1002/j.1537-2197.1960.tb07154.x.
18. ↑ Stuefer, Josef F. (1998). "Two types of division of labour in clonal plants: benefits, costs and constraints" (in en). Perspectives in Plant Ecology, Evolution and Systematics 1 (1): 47–60. doi:10.1078/1433-8319-00051. ISSN 1433-8319. Bibcode: 1998PPEES...1...47S. https://www.sciencedirect.com/science/article/abs/pii/S1433831904700049.
19. ↑ Hatfield, Jerry L.; Prueger, John H. (2015). "Temperature extremes: Effect on plant growth and development" (in en). Weather and Climate Extremes 10: 4–10. doi:10.1016/j.wace.2015.08.001. ISSN 2212-0947. Bibcode: 2015WCE....10....4H. https://www.sciencedirect.com/science/article/pii/S2212094715300116.
20. ↑ 20.0 20.1 Ahuja, I.; Fløistad, I. S.; Grande, P. O.; Tyldum, J. M.; Edvardsen, Ø M.; Steffenrem, A. (2026). "Frost hardening in seedlings of four provenances of Norway spruce in response to simulated autumn conditions with decreasing and fluctuating temperatures" (in en). IForest - Biogeosciences and Forestry 19 (2): 141. doi:10.3832/ifor5041-019. ISSN 1971-7458. https://iforest.sisef.org/contents/?id=ifor5041-019.
21. ↑ Lyons, J.M.; Raison, J.K.; Steponkus, Peter L. (1979-01-01), "The Plant Membrane in Response to Low Temperature: An Overview" (in en-US), Low Temperature Stress in Crop Plants (Academic Press): pp. 1–24, doi:10.1016/B978-0-12-460560-2.50005-4, ISBN 978-0-12-460560-2, https://www.sciencedirect.com/science/chapter/edited-volume/abs/pii/B9780124605602500054, retrieved 2026-05-05
22. ↑ 22.0 22.1 Sakai, A. (1987). Frost survival of plants : responses and adaptation to freezing stress. Internet Archive. Berlin; London : Springer. ISBN 978-3-540-17332-8. http://archive.org/details/frostsurvivalofp0000saka.
23. ↑ Pearce, R (2001). "Plant Freezing and Damage" (in en). Annals of Botany 87 (4): 417–424. doi:10.1006/anbo.2000.1352. ISSN 0305-7364. Bibcode: 2001AnBot..87..417P. https://doi.org/10.1006/anbo.2000.1352.
24. ↑ 24.0 24.1 Sakai, Akira (1979). "Freezing avoidance mechanism of primordial shoots of conifer buds1" (in en). Plant and Cell Physiology 20 (7): 1381–1390. doi:10.1093/oxfordjournals.pcp.a075937. ISSN 1471-9053. https://doi.org/10.1093/oxfordjournals.pcp.a075937.
25. ↑ Levitt, J. 1980. Responses of Plants to Environmental Stresses. Volume 1. Chilling, Freezing, and High Temperature Stresses, 2nd ed. Academic Press, New York NY. 497 p.
26. ↑ Sakai, A.; Okada, S. 1971. Freezing resistance of conifers. Silvae Genet. 20(3):91–97.
27. ↑ Michael A Dirr; Charles W Heuser, jr. (2006). "2". The Reference Manual of Woody Plant Propagation (Second ed.). Varsity Press Inc.. pp. 26, 28, 29. ISBN 0942375092.
28. ↑ Jones, Cynthia S. (1999-11-01). "An Essay on Juvenility, Phase Change, and Heteroblasty in Seed Plants". International Journal of Plant Sciences 160 (S6): –105–S111. doi:10.1086/314215. ISSN 1058-5893. PMID 10572025. Bibcode: 1999IJPlS.160S.105J.
29. ↑ Sattler, R. (1984). "Homology - a continuing challenge". Systematic Botany 9 (4): 382–394. doi:10.2307/2418787. Bibcode: 1984SysBo...9..382S.
30. ↑ Sattler, R., 1994, Homology, homeosis, and process morphology in plants. In: B.K. Hall (ed.) Homology: The hierarchical basis of comparative morphology. New York: Academic Press, pp. 423–475.
31. ↑ Kirchoff, B K; Pfeifer, E; Rutishauser, R (2008). "Plant structure ontology: How should we label plant structures with doubtful or mixed identities?". Zootaxa 1950: 103–122. doi:10.11646/zootaxa.1950.1.10.
32. ↑ Conn, Adam; Pedmale, Ullas; Chory, Joanne (2017). "High-Resolution Laser Scanning Reveals Plant Architectures that Reflect Universal Network Design Principles.". Cell Systems 5 (1): 103–122. doi:10.1016/j.cels.2017.06.017. PMID 28750198.
33. ↑ Rutishauser, R.; Isler, B. (2001). "Developmental Genetics and Morphological Evolution of Flowering Plants, Especially Bladderworts (Utricularia): Fuzzy Arberian Morphology Complements Classical Morphology". Annals of Botany 88 (6): 1173–1202. doi:10.1006/anbo.2001.1498. Bibcode: 2001AnBot..88.1173R. http://www.systbot.uzh.ch/static/personen/rolf_rutishauser_assets/Rut_Isl_AnnBot_2001.pdf.
34. ↑ James, P. J. (2009). "'Tree and Leaf': A different angle". The Linnean 25: 13–19.
35. ↑ Eckardt, NA; Baum, D (2010). "The Podostemad Puzzle: The Evolution of Unusual Morphology in the Podostemaceae". The Plant Cell 22 (7): 2131–2140. doi:10.1105/tpc.110.220711. PMID 20647343. Bibcode: 2010PlanC..22.2104E.
36. ↑ Sattler, R. (1992). "Process morphology: Structural dynamics in development and evolution". Canadian Journal of Botany 70 (4): 708–714. doi:10.1139/b92-091. Bibcode: 1992CaJB...70..708S.
37. ↑ Vergara-Silva, F. (2003). "Plants and the Conceptual Articulation of Evolutionary Developmental Biology". Biology and Philosophy 18 (2): 261–264. doi:10.1023/A:1023936102602.
38. ↑ Sattler, R. 2019. Structural and dynamic approaches to the development and evolution of plant form. In: Fusco, G. (ed) Perspectives on Evolutionary and Developmental Biology. Essays for Alessandro Minelli. Chapter 6, pp. 57–70 [1]
39. ↑ Woodger, J.H. 1967. Biological Principles. London: Routledge & Kegoan Paul (reissued with a new Introduction).
40. ↑ Jeune, B; Barabé, D; Lacroix, C (2006). "Classical and dynamic morphology: Toward a synthesis through the space of forms". Acta Biotheoretica 54 (4): 277–293. doi:10.1007/s10441-007-9007-8. PMID 17486414.
41. ↑ Minelli, A. 2018. Plant Evolutionary Developmental Biology. The Evolvability of the Phenotype. New York: Cambridge University Press.
42. ↑ Rutishauser, R. (2016). "Evolution of unusual morphologies in Lentibulariaceae (bladderworts and allies) and Podostemonaceae (river-weeds): a pictorial report at the interface of developmental biology and morphological diversification". Annals of Botany 117 (5): 811–832. doi:10.1093/aob/mcv172. PMID 26589968.
43. ↑ Rutishauser, R. (2020). "EvoDevo: Past and future of continuum and process plant morphology". Philosophies 5 (4): 166–168. doi:10.3390/philosophies5040041. PMID 5040041.
44. ↑ Sattler, R. (2024). "Morpho evo-devo of the gynoecium: Heterotopy, redefinition of the carpel and a topographic approach". Plants 13 (5): 599. doi:10.3390/plants13050599. PMID 38475445. Bibcode: 2024Plnts..13..599S.
45. ↑ Sattler, R. (2018). "Philosophy of plant morphology". Elemente der Naturwissenschaft 108: 55–79. (for an expanded version of this article see [2])
46. ↑ Kaplan, D. R., edited by Specht, C. D. 2022. Kaplan's Principles of Plant Morphology. New York: CRC Press.
47. ↑ Sattler, R. (2022). "Kaplan's Principles of Plant Morphology: A Critical Review". The Botanical Review 88 (2): 257–270. doi:10.1007/s12229-022-09280-8. Bibcode: 2022BotRv..88..257S.
48. ↑ Sattler, R.; Rutishauser, R. (2023). "Fundamentals of plant morphology and plant evo-devo (evolutionary developmental biology)". Plants 12 (1): 118–131. doi:10.3390/plants12010118. PMID 36616247.
49. ↑ Classen-Bockhoff, R. 2024. Die Pflanze: Morphologie, Entwicklung und Evolution von Vielfalt. Berlin: Springer Spektrum.
50. ↑ 50.0 50.1 Rolf Sattler (2025) Plant Evo-Devo Morphology: An Innovative New Book by Regine Claβen-Bockhoff. The Botanical Review, Dec. 8, 2025. https://doi.org/10.1007/s12229-025-09322-x
51. ↑ Rolf Sattler (2026) Articulation morphology of plants and plant evo-devo: An open morphology - empirical, dynamic, all-inclusive, and unifying. Plants Plants 15 (5), 730. https://www.mdpi.com/2223-7747/15/5/730

## External links

* Botanical Visual Glossary
* Plant morphology: fundamental issues

* v
* t
* e

Anatomy and morphology

Fields|

* Gross anatomy
* Superficial anatomy
* Neuroanatomy
* Comparative anatomy
* Transcendental anatomy
* Microscopic anatomy
* histology
* molecular

Bacteria|

* Bacterial cell structure

Protists|

* Level of organization
* Structures

Plants|

* Plant anatomy
* Plant habit
* Plant life-form/growth-form/physiognomy
* Plant morphology
* Fruit anatomy

Invertebrates|

* Body plan
* Decapod anatomy
* Gastropod anatomy
* Insect morphology
* Spider anatomy

Mammals|

* Human anatomy
* Neanderthal anatomy
* Cat anatomy
* Dog anatomy
* Horse anatomy
* Elephant anatomy
* Giraffe anatomy

Other vertebrates|

* Bird anatomy
* Fish anatomy
* Shark anatomy

Other topics|

* Allometry
* Brain morphometry
* Morphometrics
* Physiognomy

* Category
* Portal

* v
* t
* e

Botany

History of botany

Subdisciplines|

* Plant systematics
* Ethnobotany
* Paleobotany
* Plant anatomy
* Plant ecology
* Phytogeography
* Geobotany
* Flora
* Phytochemistry
* Plant pathology
* Bryology
* Phycology
* Floristics
* Dendrology
* Astrobotany

Plant groups|

* Algae
* Archaeplastida
* Bryophyte
* Non-vascular plants
* Vascular plants
* Spermatophytes
* Pteridophyte
* Gymnosperm
* Angiosperm

Plant morphology
(glossary)

| Plant cells|

* Cell wall
* Phragmoplast
* Plastid
* Plasmodesma
* Vacuole

|

Tissues|

* Meristem
* Vascular tissue
* Vascular bundle
* Ground tissue
* Mesophyll
* Cork
* Wood
* Storage organs

Vegetative|

* Root
* Rhizoid
* Bulb
* Rhizome
* Shoot
* Stem
* Leaf
* Petiole
* Cataphyll
* Bud
* Sessility

Reproductive
(Flower)|

* Flower development
* Inflorescence
* Umbel
* Raceme
* Bract
* Pedicellate
* Flower
* Aestivation
* Whorl
* Floral symmetry
* Floral diagram
* Floral formula
* Receptacle
* Hypanthium (Floral cup)
* Perianth
* Tepal
* Petal
* Sepal
* Sporophyll
* Gynoecium
* Ovary
* Ovule
* Stigma
* Archegonium
* Androecium
* Stamen
* Staminode
* Pollen
* Tapetum
* Gynandrium
* Gametophyte
* Sporophyte
* Plant embryo
* Fruit
* Fruit anatomy
* Berry
* Capsule
* Seed
* Seed dispersal
* Endosperm

Surface structures|

* Epicuticular wax
* Plant cuticle
* Epidermis
* Stoma
* Nectary
* Trichome
* Prickle

* Plant physiology
* Materials

* Nutrition
* Photosynthesis
* Chlorophyll
* Plant hormone
* Transpiration
* Turgor pressure
* Bulk flow
* Aleurone
* Phytomelanin
* Sugar
* Sap
* Starch
* Cellulose

Plant growth and habit|

* Secondary growth
* Woody plants
* Herbaceous plants
* Habit
* Vines
* Lianas
* Shrubs
* Subshrubs
* Trees
* Succulent plants

Reproduction

* Evolution
* Ecology

* Alternation of generations
* Sporangium
* Spore
* Microsporangia
* Microspore
* Megasporangium
* Megaspore
* Pollination
* Artificial
* Pollinators
* Pollen tube
* Self
* Double fertilization
* Germination
* Evolutionary development
* Evolutionary history
* timeline

Plant taxonomy|

* History of plant systematics
* Herbarium
* Biological classification
* Botanical nomenclature
* Botanical name
* Correct name
* Author citation
* International Code of Nomenclature for algae, fungi, and plants (ICN)
* International Code of Nomenclature for Cultivated Plants (ICNCP)
* Taxonomic rank
* International Association for Plant Taxonomy (IAPT)
* Plant taxonomy systems
* Cultivated plant taxonomy
* Citrus taxonomy
* cultigen
* cultivar
* Group
* grex

Practice|

* Agronomy
* Floriculture
* Forestry
* Horticulture

* Lists
* Related topics

* Botanical terms
* Botanists
* by author abbreviation
* Botanical expedition

* Category
*   * WikiProject

0.00

(0 votes)

Original source: https://en.wikipedia.org/wiki/Plant morphology. Read more

|

Retrieved from "https://handwiki.org/wiki/index.php?title=Biology:Plant_morphology&oldid=4500543"
*[v]: View this template
*[t]: Discuss this template
*[e]: Edit this template