Sieve elements are specialized cells that are important for the function of phloem, which is a highly organized tissue that transports organic compounds made during photosynthesis. Sieve elements are the major conducting cells in phloem. Conducting cells aid in transport of molecules especially for long-distance signaling. In plant anatomy, there are two main types of sieve elements. Companion cells and sieve cells originate from meristems, which are tissues that actively divide throughout a plant's lifetime. They are similar to the development of xylem, a water conducting tissue in plants whose main function is also transportation in the plant vascular system. [1] Sieve elements' major function includes transporting sugars over long distance through plants by acting as a channel. Sieve elements elongate cells containing sieve areas on their walls. Pores on sieve areas allow for cytoplasmic connections to neighboring cells, which allows for the movement of photosynthetic material and other organic molecules necessary for tissue function. Structurally, they are elongated and parallel to the organ or tissue that they are located in. Sieve elements typically lack a nucleus and contain none to a very small number of ribosomes. [2] The two types of sieve elements, sieve tube members and sieve cells, have different structures. Sieve tube members are shorter and wider with greater area for nutrient transport while sieve cells tend to be longer and narrower with smaller area for nutrient transport. Although the function of both of these kinds of sieve elements is the same, sieve cells are found in gymnosperms, non-flowering vascular plants, while sieve tube members are found in angiosperms, flowering vascular plants. [3]
Sieve elements were first discovered by the forest botanist Theodor Hartig in 1837. Since this discovery, the structure and physiology of phloem tissue has been emphasized more as there has been greater focus on its specialized components such as the sieve cells. Phloem was introduced by Carl Nägeli in 1858 after the discovery of sieve elements. Since then, multiple studies have been conducted on how sieve elements function in phloem in terms of working as a transport mechanism. [2] An example of analysis of phloem through sieve elements was conducted in the study of Arabidopsis leaves. By studying the phloem of the leaves in vivo through laser microscopy and the usage of fluorescent markers (placed in both companion cells and sieve elements), the network of companion cells with the compact sieve tubes was highlighted. The markers for sieve elements and companion cells was used to study the network and organization of phloem cells. [4]
There are two categories of sieve elements: sieve cells and sieve tube members. [5] The main functions of sieve tube members include maintaining cells and transporting necessary molecules with the help of companion cells. [6] The sieve tube members are living cells (which do not contain a nucleus) that are responsible for transporting carbohydrates throughout the plant. [7] Sieve tube members are associated with companion cells, which are cells that combine with sieve tubes to create the sieve element-companion cell complex. This allows for supply and maintenance of the plant cells and for signaling between distant organs within the plant. [6] Sieve tube members do not have ribosomes or a nucleus and thus need companion cells to help them function as transport molecules. Companion cells provide sieve tube members with proteins necessary for signaling and ATP in order to help them transfer molecules between different parts of the plant. It is the companion cells that helps transport carbohydrates from outside the cells into the sieve tube elements. [8] The companion cells also allow for bidirectional flow. [2]
While sieve tube members are responsible for a lot of the signaling necessary for the plant's organs, only some proteins are active within the sieve tubes. This is due to the fact that sieve tube members do not have ribosomes to synthesize protein as this makes it harder to determine which active proteins are specifically related to the sieve tube elements. [6]
Sieve tube members and companion cells are connected through plasmodesmata. [5] Plasmodesmata consists of channels between cell walls of adjacent plant cells for transport and cell to cell recognition. Structurally, the walls of sieve tubes tend to be dispersed with plasmodesmata grouped together and it is these areas of the tube walls and plasmodesmata that develop into sieve plates over time. Sieve tube members tend to be found largely in angiosperms. [1] They are very long and have horizontal end walls containing sieve plates. Sieve plates contain sieve pores which can regulate the size of the openings in the plates with changes in the surroundings of the plants. [3] These sieve plates are very large which means that there is a greater surface area for material transport. [5]
Sieve tube members are arranged from end to end in a longitudinal manner in order to form sieve tubes. Formed through these vertical connections between multiple sieve tube members, sieve tubes are directly responsible for the transport through the minimum resistance surrounding their walls. [8] By having the assistance of these pores that constitute a majority of the structure of sieve plates, the diameter of the sieve tubes can be regulated. This regulation is necessary for the sieve tubes to respond to changes in the environment and conditions within the organism. [5]
Sieve cells are long, conducting cells in the phloem that do not form sieve tubes. The major difference between sieve cells and sieve tube members is the lack of sieve plates in sieve cells. [1] They have a very narrow diameter and tend to be longer in length than sieve tube elements as they are generally associated with albuminous cells. [4] Similar to how Sieve Tube members are associated with companion cells, sieve cells are flanked with albuminous cells in order to aid in transporting organic material. Albuminous cells have long, unspecialized areas with ends that overlap with those of other sieve cells and contain nutrients and store food in order to nourish tissues. [7] They enable the sieve cells to be connected to parenchyma, functional tissue in the organs, which helps to stabilize the tissue and transport nutrients. Sieve cells are also associated with gymnosperms because they lack the companion cell and sieve member complexes that angiosperms have. [9] Sieve cells are very uniform and have an even distribution across of sieve areas. Their narrow pores are necessary in their function in most seedless vascular plants and gymnosperms which lack sieve-tube members and only have sieve cells to transport molecules. [1] While sieve cells have smaller sieve areas, they are still distributed across several cells to still effectively transport material to various tissue within the plant. [2]
Sieve cell associated albuminous cells work between phloem and parenchyma. They connect parenchyma with mature sieve cells to help participate in transport of cells. There can be many of these albuminous cells that belong to one sieve cell, depending on the function of the tissue or organ. [1]
Sieve pores are very common in the areas that have overlapping sieve cells. Callose levels are measure in order to observe the activity of sieve cells. Callose acts as a block to the sieve pores that are present in both of these sieve elements. A lack callose suggests that the sieve elements are more active and therefore can regulate their pores more actively in response to environmental changes. [10]
Because the plant vascular system is vital in growth and development of plant cells and the organs within the plant, the role of sieve elements in the transport of necessary carbohydrates and macromolecules is largely expanded. This can be applied to agriculture to observe the way resources are distributed to various parts of the plant. Plasmodesmata connect companion cells to sieve elements and parenchyma cells can connect the sieve tubes to various tissues within the plant. This system between the plasmodesmata, companion cells, and sieve tubes allow for the delivery of necessary metabolites. The yield of agricultural product could potentially be increased to maximize the delivery system of these specialized cells within the phloem in a way that diffusion can be maximized. It has been discovered that the angiosperm phloem can use the sieve tubes as a way to transport various forms of RNA to sink tissues which can help alter transcriptional activity. Sink tissues are tissues that are in the process of growth and need nutrients. Having Sieve elements transport additional nutrients to sink tissues can speed up the growth process, which can affect plant growth and development. Over time, rapid growth has the potential of leading to greater agricultural output. [11]
A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.
Plant cells are the cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capability to perform photosynthesis and store starch, a large vacuole that regulates turgor pressure, the absence of flagella or centrioles, except in the gametes, and a unique method of cell division involving the formation of a cell plate or phragmoplast that separates the new daughter cells.
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of the xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word xylem is derived from the Ancient Greek word ξύλον (xylon), meaning "wood"; the best-known xylem tissue is wood, though it is found throughout a plant. The term was introduced by Carl Nägeli in 1858.
Phloem is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as photosynthates, in particular the sugar sucrose, to the rest of the plant. This transport process is called translocation. In trees, the phloem is the innermost layer of the bark, hence the name, derived from the Ancient Greek word φλοιός (phloiós), meaning "bark". The term was introduced by Carl Nägeli in 1858.
Vascular plants, also called tracheophytes or collectively Tracheophyta, form a large group of land plants that have lignified tissues for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue to conduct products of photosynthesis. Vascular plants include the clubmosses, horsetails, ferns, gymnosperms, and angiosperms. Scientific names for the group include Tracheophyta, Tracheobionta and Equisetopsida sensu lato. Some early land plants had less developed vascular tissue; the term eutracheophyte has been used for all other vascular plants, including all living ones.
In biology, tissue is a historically derived biological organizational level between cells and a complete organ. A tissue is therefore often thought of as an assembly of similar cells and their extracellular matrix from the same embryonic origin that together carry out a specific function. Organs are then formed by the functional grouping together of multiple tissues.
The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xylem inwards, towards the pith, and secondary phloem outwards, towards the bark.
A tracheid is a long and tapered lignified cell in the xylem of vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms use another type of conductive cell, called vessel elements, to transport water through the xylem. The main functions of tracheid cells are to transport water and inorganic salts, and to provide structural support for trees. There are often pits on the cell walls of tracheids, which allows for water flow between cells. Tracheids are dead at functional maturity and do not have a protoplast. The wood (softwood) of gymnosperms such as pines and other conifers is mainly composed of tracheids. Tracheids are also the main conductive cells in the primary xylem of ferns.
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the integument, forming its outer layer, the nucellus, and the female gametophyte in its center. The female gametophyte — specifically termed a megagametophyte— is also called the embryo sac in angiosperms. The megagametophyte produces an egg cell for the purpose of fertilization. The ovule is a small structure present in the ovary. It is attached to the placenta by a stalk called a funicle. The funicle provides nourishment to the ovule.
Sap is a fluid transported in xylem cells or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant.
Sclereids are a reduced form of sclerenchyma cells with highly thickened, lignified cellular walls that form small bundles of durable layers of tissue in most plants. The presence of numerous sclereids form the cores of apples and produce the gritty texture of guavas.
The pericycle is a cylinder of parenchyma or sclerenchyma cells that lies just inside the endodermis and is the outer most part of the stele of plants.
Plasmodesmata are microscopic channels which traverse the cell walls of plant cells and some algal cells, enabling transport and communication between them. Plasmodesmata evolved independently in several lineages, and species that have these structures include members of the Charophyceae, Charales, Coleochaetales and Phaeophyceae, as well as all embryophytes, better known as land plants. Unlike animal cells, almost every plant cell is surrounded by a polysaccharide cell wall. Neighbouring plant cells are therefore separated by a pair of cell walls and the intervening middle lamella, forming an extracellular domain known as the apoplast. Although cell walls are permeable to small soluble proteins and other solutes, plasmodesmata enable direct, regulated, symplastic transport of substances between cells. There are two forms of plasmodesmata: primary plasmodesmata, which are formed during cell division, and secondary plasmodesmata, which can form between mature cells.
A vessel element or vessel member is one of the cell types found in xylem, the water conducting tissue of plants. Vessel elements are found in angiosperms but absent from gymnosperms such as conifers. Vessel elements are the main feature distinguishing the "hardwood" of angiosperms from the "softwood" of conifers.
The Hernandiaceae are a family of flowering plants (angiosperms) in the order Laurales. Consisting of five genera with about 58 known species, they are distributed over the world's tropical areas, some of them widely distributed in coastal areas, but they occur from sea level to over 2000 m.
The ascent of sap in the xylem tissue of plants is the upward movement of water and minerals from the root to the aerial parts of the plant. The conducting cells in xylem are typically non-living and include, in various groups of plants, vessel members and tracheids. Both of these cell types have thick, lignified secondary cell walls and are dead at maturity. Although several mechanisms have been proposed to explain how sap moves through the xylem, the cohesion-tension mechanism has the most support. Although cohesion-tension has received criticism due to the apparent existence of large negative pressures in some living plants, experimental and observational data favor this mechanism.
Callose is a plant polysaccharide. Its production is due to the glucan synthase-like gene (GLS) in various places within a plant. It is produced to act as a temporary cell wall in response to stimuli such as stress or damage. Callose is composed of glucose residues linked together through β-1,3-linkages, and is termed a β-glucan. It is thought to be manufactured at the cell wall by callose synthases and is degraded by β-1,3-glucanases. Callose is very important for the permeability of plasmodesmata (Pd) in plants; the plant's permeability is regulated by plasmodesmata callose (PDC). PDC is made by callose synthases and broken down by β-1,3-glucanases (BGs). The amount of callose that is built up at the plasmodesmatal neck, which is brought about by the interference of callose synthases (CalSs) and β-1,3-glucanases, determines the conductivity of the plasmodesmata.
The pressure flow hypothesis, also known as the mass flow hypothesis, is the best-supported theory to explain the movement of sap through the phloem. It was proposed by Ernst Münch, a German plant physiologist in 1930. A high concentration of organic substances, particularly sugar, inside cells of the phloem at a source, such as a leaf, creates a diffusion gradient that draws water into the cells from the adjacent xylem. This creates turgor pressure, also known as hydrostatic pressure, in the phloem. Movement of phloem sap occurs by bulk flow from sugar sources to sugar sinks. The movement in phloem is bidirectional, whereas, in xylem cells, it is unidirectional (upward). Because of this multi-directional flow, coupled with the fact that sap cannot move with ease between adjacent sieve-tubes, it is not unusual for sap in adjacent sieve-tubes to be flowing in opposite directions.
A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, photosynthesis takes place here, stores nutrients, and produces new living tissue. The stem can also be called halm or haulm or culms.
Plants are constantly exposed to different stresses that result in wounding. Plants have adapted to defend themselves against wounding events, like herbivore attacks or environmental stresses. There are many defense mechanisms that plants rely on to help fight off pathogens and subsequent infections. Wounding responses can be local, like the deposition of callose, and others are systemic, which involve a variety of hormones like jasmonic acid and abscisic acid.