Phylloclades and cladodes are flattened,

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

shoots, which are usually considered to be modified

branch A branch, also called a ramus in botany, is a stem that grows off from another stem, or when structures like veins in leaves are divided into smaller veins. History and etymology In Old English, there are numerous words for branch, includ ...

es. The two terms are used either differently or interchangeably by different authors. ''

Phyllocladus ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually placed in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...

'', a genus of conifer, is named after these structures. Phylloclades/cladodes have been identified in fossils dating from as early as the

Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...

Definition and morphology

The term "phylloclade" is from the

Neo-Latin Neo-LatinSidwell, Keith ''Classical Latin-Medieval Latin-Neo Latin'' in ; others, throughout. (also known as New Latin and Modern Latin) is the style of written Latin used in original literary, scholarly, and scientific works, first in Italy d ...

''phyllocladium'', itself derived from

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

''phyllo'', leaf, and ''klados'', branch. Definitions of the terms "phylloclade" and "cladode" vary. All agree that they are flattened structures that are photosynthetic and resemble leaf-like branches. In one definition, phylloclades are a subset of cladodes, namely those that greatly resemble or perform the function of

leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...

, as in Butcher's broom (''Ruscus aculeatus'') as well as ''

Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University ...

'' and some ''

Asparagus Asparagus (''Asparagus officinalis'') is a perennial flowering plant species in the genus ''Asparagus (genus), Asparagus'' native to Eurasia. Widely cultivated as a vegetable crop, its young shoots are used as a spring vegetable. Description ...

'' species. By an alternative definition, cladodes are distinguished by their limited growth and that they involve only one or two internodes. By this definition, some of the most leaf-like structures are cladodes, rather than phylloclades. By that definition, ''Phyllanthus'' has phylloclades, but ''Ruscus'' and ''Asparagus'' have cladodes. Another definition uses "phylloclade" to refer a portion of a leaf-like stem or branch with multiple nodes and internodes, and "cladode" for a single internode of a phylloclade. Although phylloclades are usually interpreted as modified branches, developmental studies have shown that they are intermediate between leaves and branches as their name indicates. Molecular genetic investigations have confirmed these findings. For example, Hirayama et al. (2007) showed that the phylloclade of ''Ruscus aculeatus'' "is not homologous to either the shoot or the leaf, but that it has a double organ identity," which means that it combines shoot and leaf processes.

Similar structures

* Aristate leaves end in a stiff point that may continue the primary leaf vein; this can resemble the stem end of a phylloclade/cladode. * Epiphylly: flowers and fruit develop "on a leaf". A stem and a leaf are merged with one another. Examples include ''Monophyllaea'' in family ''

Gesneriaceae Gesneriaceae, the gesneriad family, is a family (biology), family of flowering plants consisting of about 152 genera and ca. 3,540 species in the tropics and subtropics of the Old World (almost all Didymocarpoideae) and the New World (most Ges ...

'' and '' Helwingia'' in

Helwingiaceae The genus ''Helwingia'' consists of shrubs or rarely small trees native to eastern Asia, the Himalayas, and northern Indochina. It is the only genus in the family Helwingiaceae. Description The plants have alternate, evergreen or deciduous leave ...

Illustrations

image:Illustration Ruscus aculeatus0.jpg, Botanical illustration of ''Ruscus aculeatus'' showing leaf-like phylloclades/cladodes image:RuscusPhylloclade.jpg, Phylloclade/cladode of ''Ruscus'' sp. showing the spine formed by the stem axis image:Starr 031114-0007 Asparagus asparagoides.jpg, Leaf-like cladodes/phylloclades of ''

Asparagus asparagoides ''Asparagus asparagoides'', commonly known as bridal creeper, bridal-veil creeper, ''gnarboola'', smilax or smilax asparagus, is a herbaceous climbing plant of the family Asparagaceae native to eastern and southern Africa. Sometimes grown as an ...

'' image:Helwingia japonica1.jpg, Epiphylly in '' Helwingia japonica'' for comparison

References

{{Commons category, Cladodes, Phylloclade Plant morphology Plant anatomy