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Phytomorphology is the study of the physical form and external structure of
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s.Raven, P. H., R. F. Evert, & S. E. Eichhorn. ''Biology of Plants'', 7th ed., page 9. (New York: W. H. Freeman, 2005). . This is usually considered distinct from
plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plan ...
, which is the study of the internal
structure A structure is an arrangement and organization of interrelated elements in a material object or system, or the object or system so organized. Material structures include man-made objects such as buildings and machines and natural objects such a ...
of plants, especially at the microscopic level. Plant morphology is useful in the visual identification of plants. Recent studies in
molecular biology Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and phys ...
started to investigate the molecular processes involved in determining the conservation and diversification of plant morphologies. In these studies
transcriptome The transcriptome is the set of all RNA transcripts, including coding and non-coding, in an individual or a population of cells. The term can also sometimes be used to refer to all RNAs, or just mRNA, depending on the particular experiment. The t ...
conservation patterns were found to mark crucial
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the s ...
transitions during the
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
life cycle which may result in evolutionary constraints limiting diversification.


Scope

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". There are four major areas of investigation in plant morphology, and each overlaps with another field of the
biological sciences Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
. First of all, morphology is
comparative general linguistics, the comparative is a syntactic construction that serves to express a comparison between two (or more) entities or groups of entities in quality or degree - see also comparison (grammar) for an overview of comparison, as well ...
, 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 A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family (biology), family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanic ...
, oak, and
cabbage Cabbage, comprising several cultivars of ''Brassica oleracea'', is a leafy green, red (purple), or white (pale green) biennial plant grown as an annual vegetable crop for its dense-leaved heads. It is descended from the wild cabbage ( ''B.&n ...
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 A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Gree ...
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 Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
and
paleobotany Paleobotany, which is also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeog ...
. Secondly, plant morphology observes both the vegetative (somatic) structures of plants, as well as the reproductive structures. The vegetative structures of
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s includes the study of the shoot system, composed of stems and
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
, as well as the
root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
system. The reproductive structures are more varied, and are usually specific to a particular group of plants, such as
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s and
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s, fern sori, and
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ...
capsules. The detailed study of reproductive structures in plants led to the discovery of the
alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of life cycle in plants and algae. It consists of a multicellular haploid sexual phase, the gametophyte, which has a single set of chromosomes alt ...
found in all plants and most
alga Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
e. This area of plant morphology overlaps with the study of
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
and plant systematics. 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 An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
, and
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living an ...
, the study of cells using
optical microscopy Optics is the branch of physics that studies the behaviour and properties of light, including its interactions with matter and the construction of instruments that use or detect it. Optics usually describes the behaviour of visible, ultravio ...
. At this scale, plant morphology overlaps with
plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plan ...
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 In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
will vary from species to species, as will the appearance of a plant as a
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
,
herb In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicina ...
, or
grass Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns a ...
. 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 Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry (b ...
and
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
.


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: # Homology - the structure is similar between the two species because of shared ancestry and common genetics. # Convergence - the structure is similar between the two species because of independent adaptation to common environmental pressures. Understanding which characteristics and structures belong to each type is an important part of understanding plant
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. The evolutionary biologist relies on the plant morphologist to interpret structures, and in turn provides
phylogenies A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of plant relationships that may lead to new morphological insights.


Homology

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.


Convergence

When structures in different species are believed to exist and develop as a result of common adaptive responses to environmental pressure, those structures are termed ''convergent''. For example, the fronds of '' Bryopsis plumosa'' and stems of '' Asparagus setaceus'' both have the same feathery branching appearance, even though one is an alga and one is a flowering plant. The similarity in overall structure occurs independently as a result of convergence. The growth form of many
cacti A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Gree ...
and species of ''
Euphorbia ''Euphorbia'' is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to t ...
'' is very similar, even though they belong to widely distant families. The similarity results from common solutions to the problem of surviving in a hot, dry environment.


Vegetative and reproductive characteristics

Plant morphology treats both the vegetative structures of plants, as well as the reproductive structures. The vegetative (somatic) structures of
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s include two major organ systems: (1) a shoot system, composed of stems and
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
, 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
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s and
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
s are only found in the
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
s; sori are only found in
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s; and seed cones are only found in
conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
s and other gymnosperms. Reproductive characters are therefore regarded as more useful for the classification of plants than vegetative characters.


Use in identification

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. # Quantitative characters are morphological features that can be counted or measured for example a plant species has flower petals 10–12 mm wide. # 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

The detailed study of reproductive structures in plants led to the discovery of the
alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of life cycle in plants and algae. It consists of a multicellular haploid sexual phase, the gametophyte, which has a single set of chromosomes alt ...
, found in all plants and most
alga Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
e, by the German botanist Wilhelm Hofmeister. This discovery is one of the most important made in all of plant morphology, since it provides a common basis for understanding the life cycle of all plants.


Pigmentation in plants

The primary function of pigments in plants is
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
, which uses the green pigment
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
along with several red and yellow pigments that help to capture as much light energy as possible. Pigments are also an important factor in attracting
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s to flowers to encourage
pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...
. Plant pigments include a variety of different kinds of molecule, including porphyrins,
carotenoid Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, ...
s,
anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...
s and betalains. All biological pigments selectively absorb certain wavelengths of
light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 t ...
while reflecting others. The light that is absorbed may be used by the plant to power chemical reactions, while the reflected wavelengths of light determine the color the pigment will appear to the eye.


Morphology in development

Plant development is the process by which structures originate and mature as a plant grows. It is a subject studies in
plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plan ...
and
plant physiology Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry (b ...
as well as plant morphology. The process of development in plants is fundamentally different from that seen in
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
animals. When an animal
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
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 located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. The properties of organization seen in a plant are emergent properties which are more than the sum of the individual parts. "The assembly of these tissues and functions into an integrated multicellular organism yields not only the characteristics of the separate parts and processes but also quite a new set of characteristics which would not have been predictable on the basis of examination of the separate parts." In other words, knowing everything about the molecules in a plant are not enough to predict characteristics of the cells; and knowing all the properties of the cells will not predict all the properties of a plant's structure.


Growth

A
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
begins from a single celled
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicell ...
, formed by
fertilisation Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
of an egg cell by a sperm cell. From that point, it begins to divide to form a plant
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
through the process of embryogenesis. As this happens, the resulting cells will organize so that one end becomes the first root, while the other end forms the tip of the shoot. In
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
plants, the embryo will develop one or more "seed leaves" (
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...
s). By the end of embryogenesis, the young plant will have all the parts necessary to begin in its life. 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. Branching occurs when small clumps of cells left behind by the meristem, and which have not yet undergone
cellular differentiation Cellular differentiation is the process in which a stem cell alters from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellular ...
to form a specialized tissue, begin to grow as 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 In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...
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. 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 Phototropism is the growth of an organism in response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the light contain a hor ...
), gravity ( gravitropism), water, ( hydrotropism), and physical contact ( thigmotropism). Plant growth and development are mediated by specific
plant hormone Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pat ...
s and plant growth regulators (PGRs) (Ross et al. 1983).Ross, S.D.; Pharis, R.P.; Binder, W.D. 1983. Growth regulators and conifers: their physiology and potential uses in forestry. p. 35–78 ''in'' Nickell, L.G. (Ed.), Plant growth regulating chemicals. Vol. 2, CRC Press, Boca Raton FL. 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.


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.


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.


Positional effects

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.


Environmental effects

The way in which new structures mature as they are produced may be affected by the point in the plants life when they begin to develop, as well as by the environment to which the structures are exposed. This can be seen in
aquatic plant Aquatic plants are plants that have adapted to living in aquatic environments ( saltwater or freshwater). They are also referred to as hydrophytes or macrophytes to distinguish them from algae and other microphytes. A macrophyte is a plant that ...
s and emergent plants.


=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. 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.Glerum, C. 1985. Frost hardiness of coniferous seedlings: principles and applications. p. 107–123 ''in ''Duryea, M.L. (Ed.). Proceedings: Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Workshop, October 1984, Oregon State Univ., For. Res. Lab., Corvallis OR. Intracellular freezing usually kills the cell regardless of the hardiness of the plant and its tissues.Lyons, J.M.; Raison, J.K.; Steponkus, P.L. 1979. The plant membrane in response to low temperature: an overview. p. 1–24 ''in'' Lyons, J.M.; Graham, D.; Raison, J.K. (Eds.). Low Temperature Stress in Crop Plants. Academic Press, New York NY. 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"Sakai, A.; Larcher, W. (Eds.) 1987. Frost Survival of Plants. Springer-Verlag. 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. 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. The rate of cooling has been shown to influence the frost resistance of tissues,Sakai, A. 1979a. Freezing avoidance mechanism of primordial shoots of conifer buds. Plant Cell Physiol. 20:1381–1390. 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.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. Sakai (1979a) 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 Liquid nitrogen—LN2—is nitrogen in a liquid state at low temperature. Liquid nitrogen has a boiling point of about . It is produced industrially by fractional distillation of liquid air. It is a colorless, low viscosity liquid that is wid ...
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. The hardiness of the winter buds of such
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
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,Sakai, A.; Okada, S. 1971. Freezing resistance of conifers. Silvae Genet. 20(3):91–97. given similar states of dormancy.


Juvenility

The organs and tissues produced by a young plant, such as a
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (emb ...
, 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 recognize 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. The transition from early to late growth forms is referred to as ' vegetative phase change', but there is some disagreement about terminology.


Modern Innovations

Rolf Sattler has revised fundamental concepts of comparative morphology such as the concept of homology. He emphasized that homology should also include partial homology and quantitative homology. 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. A recent study conducted by Stalk 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 optimizes 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." 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 Fuzzy logic is a form of many-valued logic in which the truth value of variables may be any real number between 0 and 1. It is employed to handle the concept of partial truth, where the truth value may range between completely true and completel ...
, “Arberian” to
Agnes Arber Agnes Robertson Arber FRS (23 February 1879 – 22 March 1960) was a British plant morphologist and anatomist, historian of botany and philosopher of biology. She was born in London but lived most of her life in Cambridge, including the las ...
. Rutishauser and Isler emphasized that this approach is not only supported by many morphological data but also by evidence from molecular genetics. More recent evidence from molecular genetics provides further support for continuum morphology. James (2009) concluded that "it is now widely accepted that... radiality haracteristic of most stemsand dorsiventrality haracteristic of leavesare but extremes of a continuous spectrum. In fact, it is simply the timing of the KNOX gene expression!." Eckardt and Baum (2010) concluded that "it is now generally accepted that compound leaves express both leaf and shoot properties.” Process morphology describes and analyzes the dynamic continuum of plant form. According to this approach, structures do not ''have'' process(es), they ''are'' process(es). Thus, the structure/process dichotomy is overcome by "an enlargement of our concept of 'structure' so as to include and recognize 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". 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). 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. 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. Rutishauser (2020) discussed the past and future of plant evo-devo. 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. One important and unique event in plant morphology of the 21st century was the publication of Kaplan's Principles of Plant Morphology by Donald R. Kaplan, edited by Chelsea D. Specht (2020). 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 all modern conceptional innovations.


See also

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Glossary of plant morphology This page provides a glossary of plant morphology. Botanists and other biologists who study plant morphology use a number of different terms to classify and identify plant organs and parts that can be observed using no more than a handheld magnif ...
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Plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plan ...
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Plant identification In biology, determination is the process of matching a specimen of an organism to a known taxon, for example identifying a plant. The term is also used in cellular biology, where it means the act of the differentiation of stem cells becoming fix ...
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Plant physiology Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry (b ...
* Plant evolutionary developmental biology *
Taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...


References


External links


Botanical Visual Glossary


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