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119-476: Nectar is a viscous , sugar -rich liquid produced by plants in glands called nectaries , either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries , which provide a nutrient source to animal mutualists , which in turn provide herbivore protection . Common nectar-consuming pollinators include mosquitoes , hoverflies , wasps , bees , butterflies and moths , hummingbirds , honeyeaters and bats . Nectar plays

238-435: A magnetic field , possibly to the point of behaving like a solid. The viscous forces that arise during fluid flow are distinct from the elastic forces that occur in a solid in response to shear, compression, or extension stresses. While in the latter the stress is proportional to the amount of shear deformation, in a fluid it is proportional to the rate of deformation over time. For this reason, James Clerk Maxwell used

357-416: A nutrient -rich tissue , inside the seed . The two central-cell maternal nuclei (polar nuclei) that contribute to the endosperm arise by mitosis from the single meiotic product that also gave rise to the egg. Therefore, maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo. One primitive species of flowering plant, Nuphar polysepala , has endosperm that

476-436: A constant viscosity ( non-Newtonian fluids ) cannot be described by a single number. Non-Newtonian fluids exhibit a variety of different correlations between shear stress and shear rate. One of the most common instruments for measuring kinematic viscosity is the glass capillary viscometer. In coating industries, viscosity may be measured with a cup in which the efflux time is measured. There are several sorts of cup—such as

595-521: A crucial role in the foraging economics and evolution of nectar-eating species; for example, nectar foraging behavior is largely responsible for the divergent evolution of the African honey bee, A. m. scutellata and the western honey bee . Nectar is an economically important substance as it is the sugar source for honey . It is also useful in agriculture and horticulture because the adult stages of some predatory insects feed on nectar. For example,

714-407: A few days after; therefore, in most mammals, it is more common for ejaculation to precede ovulation than vice versa. When sperm are deposited into the anterior vagina, they are not capable of fertilisation (i.e., non-capacitated) and are characterised by slow linear motility patterns. This motility, combined with muscular contractions enables sperm transport towards the uterus and oviducts . There

833-506: A fluid, just as thermal conductivity characterizes heat transport, and (mass) diffusivity characterizes mass transport. This perspective is implicit in Newton's law of viscosity, τ = μ ( ∂ u / ∂ y ) {\displaystyle \tau =\mu (\partial u/\partial y)} , because the shear stress τ {\displaystyle \tau } has units equivalent to

952-415: A momentum flux , i.e., momentum per unit time per unit area. Thus, τ {\displaystyle \tau } can be interpreted as specifying the flow of momentum in the y {\displaystyle y} direction from one fluid layer to the next. Per Newton's law of viscosity, this momentum flow occurs across a velocity gradient, and the magnitude of the corresponding momentum flux

1071-420: A more violent and rapid non-linear motility pattern as sperm approach the oocyte. The capacitated spermatozoon and the oocyte meet and interact in the ampulla of the fallopian tube . Rheotaxis, thermotaxis and chemotaxis are known mechanisms that guide sperm towards the egg during the final stage of sperm migration. Spermatozoa respond (see Sperm thermotaxis ) to the temperature gradient of ~2 °C between

1190-403: A nocturnal scent from the labellum epidermis. Ophrys labella have dome-shaped, papillate, dark-staining epidermal cells forming osmophores. Narcissus emit pollinator specific volatiles from the corona . Viscous Viscosity quantifies the internal frictional force between adjacent layers of fluid that are in relative motion. For instance, when a viscous fluid is forced through

1309-444: A number of predacious or parasitoid wasps (e.g., the social wasp species Apoica flavissima ) rely on nectar as a primary food source. In turn, these wasps then hunt agricultural pest insects as food for their young. Nectar is most often associated with flowering plants angiosperms , but it is also produced by other groups, including ferns . Nectar is derived from Greek νέκταρ , the fabled drink of eternal life. Some derive

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1428-613: A preference for plants with extrafloral nectaries, particularly some species of ants and wasps , which have been observed to defend the plants bearing them. Acacia is one example of a plant whose nectaries attract ants, which protect the plant from other insect herbivores . Among passion flowers , for example, extrafloral nectaries prevent herbivores by attracting ants and deterring two species of butterflies from laying eggs. In many carnivorous plants , extrafloral nectaries are also used to attract insect prey . Charles Darwin understood that extrafloral nectar "though small in quantity,

1547-461: A protective function. The Nicotiana attenuata , a tobacco plant native to the US state of Utah , uses several volatile aromas to attract pollinating birds and moths. The strongest such aroma is benzylacetone , but the plant also adds bitter nicotine , which is less aromatic, so may not be detected by the bird until after taking a drink. Researchers speculate the purpose of this addition is to discourage

1666-517: A short period lasting some days; a queen may mate with eight or more drones . She then stores the sperm for the rest of her life, perhaps for five years or more. In many fungi (except chytrids ), as in some protists, fertilisation is a two step process. First, the cytoplasms of the two gamete cells fuse (called plasmogamy ), producing a dikaryotic or heterokaryotic cell with multiple nuclei. This cell may then divide to produce dikaryotic or heterokaryotic hyphae . The second step of fertilisation

1785-465: A signal from the female gametophyte. Specific proteins called FER protein kinases present in the ovule control the production of highly reactive derivatives of oxygen called reactive oxygen species (ROS). ROS levels have been shown via GFP to be at their highest during floral stages when the ovule is the most receptive to pollen tubes, and lowest during times of development and following fertilisation. High amounts of ROS activate Calcium ion channels in

1904-416: A specific fluid state. To standardize comparisons among experiments and theoretical models, viscosity data is sometimes extrapolated to ideal limiting cases, such as the zero shear limit, or (for gases) the zero density limit. Transport theory provides an alternative interpretation of viscosity in terms of momentum transport: viscosity is the material property which characterizes momentum transport within

2023-401: A spring is proportional to the distance displaced from equilibrium. Stresses which can be attributed to the deformation of a material from some rest state are called elastic stresses. In other materials, stresses are present which can be attributed to the deformation rate over time . These are called viscous stresses. For instance, in a fluid such as water the stresses which arise from shearing

2142-567: A subject of semantic arguments about the beginning of pregnancy , typically in the context of the abortion debate. Upon gastrulation , which occurs around 16 days after fertilisation, the implanted blastocyst develops three germ layers, the endoderm, the ectoderm and the mesoderm, and the genetic code of the father becomes fully involved in the development of the embryo; later twinning is impossible. Additionally, interspecies hybrids survive only until gastrulation and cannot further develop. However, some human developmental biology literature refers to

2261-478: A thick layer of extracellular matrix that surrounds the egg and is similar to the role of the vitelline membrane in sea urchins, binds the sperm. Unlike sea urchins, the sperm binds to the egg before the acrosomal reaction. ZP3 , a glycoprotein in the zona pellucida, is responsible for egg/sperm adhesion in humans. The receptor galactosyltransferase (GalT) binds to the N-acetylglucosamine residues on

2380-400: A thin spike at the head of the sperm called the acrosomal process . The sperm binds to the egg through another ligand reaction between receptors on the vitelline membrane . The sperm surface protein bindin, binds to a receptor on the vitelline membrane identified as EBR1. Fusion of the plasma membranes of the sperm and egg are likely mediated by bindin. At the site of contact, fusion causes

2499-427: A tiny pore called a micropyle . The sperm are transferred from the pollen through the pollen tube to the ovule where the egg is fertilised. In flowering plants , two sperm cells are released from the pollen tube, and a second fertilisation event occurs involving the second sperm cell and the central cell of the ovule, which is a second female gamete. Unlike animal sperm which is motile, the sperm of most seed plants

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2618-416: A tube, it flows more quickly near the tube's center line than near its walls. Experiments show that some stress (such as a pressure difference between the two ends of the tube) is needed to sustain the flow. This is because a force is required to overcome the friction between the layers of the fluid which are in relative motion. For a tube with a constant rate of flow, the strength of the compensating force

2737-448: Is karyogamy , the fusion of the nuclei to form a diploid zygote. In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants. There are three types of fertilisation processes in protozoa: Algae, like some land plants, undergo alternation of generations . Some algae are isomorphic, where both the sporophyte (2n) and gameteophyte (n) are the same morphologically. When algae reproduction

2856-664: Is 1 cP divided by 1000 kg/m^3, close to the density of water. The kinematic viscosity of water at 20 °C is about 1 cSt. The most frequently used systems of US customary, or Imperial , units are the British Gravitational (BG) and English Engineering (EE). In the BG system, dynamic viscosity has units of pound -seconds per square foot (lb·s/ft ), and in the EE system it has units of pound-force -seconds per square foot (lbf·s/ft ). The pound and pound-force are equivalent;

2975-457: Is a linear combination of the shear and bulk viscosities that describes the reaction of a solid elastic material to elongation. It is widely used for characterizing polymers. In geology , earth materials that exhibit viscous deformation at least three orders of magnitude greater than their elastic deformation are sometimes called rheids . Viscosity is measured with various types of viscometers and rheometers . Close temperature control of

3094-462: Is a calculation derived from tests performed on drilling fluid used in oil or gas well development. These calculations and tests help engineers develop and maintain the properties of the drilling fluid to the specifications required. Nanoviscosity (viscosity sensed by nanoprobes) can be measured by fluorescence correlation spectroscopy . The SI unit of dynamic viscosity is the newton -second per square meter (N·s/m ), also frequently expressed in

3213-448: Is a pH gradient within the micro-environment of the female reproductive tract such that the pH near the vaginal opening is lower (approximately 5) than the oviducts (approximately 8). The sperm-specific pH-sensitive calcium transport protein called CatSper increases the sperm cell permeability to calcium as it moves further into the reproductive tract. Intracellular calcium influx contributes to sperm capacitation and hyperactivation, causing

3332-550: Is a viscosity tensor that maps the velocity gradient tensor ∂ v k / ∂ r ℓ {\displaystyle \partial v_{k}/\partial r_{\ell }} onto the viscous stress tensor τ i j {\displaystyle \tau _{ij}} . Since the indices in this expression can vary from 1 to 3, there are 81 "viscosity coefficients" μ i j k l {\displaystyle \mu _{ijkl}} in total. However, assuming that

3451-610: Is also estimated that about 42% of flowering plants exhibit a mixed mating system in nature. In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-fertilised, outcrossed or a mixture of progeny types. The transition from cross-fertilisation to self-fertilisation is the most common evolutionary transition in plants, and has occurred repeatedly in many independent lineages. About 10-15% of flowering plants are predominantly self-fertilising. Under circumstances where pollinators or mates are rare, self-fertilisation offers

3570-532: Is called the rate of shear deformation or shear velocity , and is the derivative of the fluid speed in the direction parallel to the normal vector of the plates (see illustrations to the right). If the velocity does not vary linearly with y {\displaystyle y} , then the appropriate generalization is: where τ = F / A {\displaystyle \tau =F/A} , and ∂ u / ∂ y {\displaystyle \partial u/\partial y}

3689-402: Is described as oogamous, the male and female gametes are different morphologically, where there is a large non-motile egg for female gametes, and the male gamete are uniflagellate (motile). Via the process of syngamy, these will form a new zygote, regenerating the sporophyte generation again. Meiosis results in a random segregation of the genes that each parent contributes. Each parent organism

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3808-448: Is determined by the viscosity. The analogy with heat and mass transfer can be made explicit. Just as heat flows from high temperature to low temperature and mass flows from high density to low density, momentum flows from high velocity to low velocity. These behaviors are all described by compact expressions, called constitutive relations , whose one-dimensional forms are given here: where ρ {\displaystyle \rho }

3927-405: Is diploid, resulting from the fusion of a sperm with one, rather than two, maternal nuclei. It is believed that early in the development of angiosperm lineages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio. In many plants, the development of the flesh of the fruit

4046-540: Is equal to the SI millipascal second (mPa·s). The SI unit of kinematic viscosity is square meter per second (m /s), whereas the CGS unit for kinematic viscosity is the stokes (St, or cm ·s = 0.0001 m ·s ), named after Sir George Gabriel Stokes . In U.S. usage, stoke is sometimes used as the singular form. The submultiple centistokes (cSt) is often used instead, 1 cSt = 1 mm ·s  = 10  m ·s . 1 cSt

4165-910: Is greedily sought by insects" but believed that "their visits do not in any way benefit the plant". Instead, he believed that extrafloral nectaries were excretory in nature ( hydathodes ). Their defensive functions were first recognized by the Italian botanist Federico Delpino in his important monograph Funzione mirmecofila nel regno vegetale (1886). Delpino's study was inspired by a disagreement with Darwin, with whom he corresponded regularly. Extrafloral nectaries have been reported in over 3941 species of vascular plants belonging to 745 genera and 108 families , 99.7% of which belong to flowering plants (angiosperms), comprising 1.0 to 1.8% of all known species. They are most common among eudicots , occurring in 3642 species (of 654 genera and 89 families), particularly among rosids which comprise more than half of

4284-419: Is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of the style before reaching the ovary. Then near the receptacle, it breaks through the ovule through the micropyle (an opening in the ovule wall) and the pollen tube "bursts" into the embryo sac, releasing sperm. The growth of

4403-576: Is in terms of the standard (scalar) viscosity μ {\displaystyle \mu } and the bulk viscosity κ {\displaystyle \kappa } such that α = κ − 2 3 μ {\displaystyle \alpha =\kappa -{\tfrac {2}{3}}\mu } and β = γ = μ {\displaystyle \beta =\gamma =\mu } . In vector notation this appears as: where δ {\displaystyle \mathbf {\delta } }

4522-658: Is not a fundamental law of nature, but rather a constitutive equation (like Hooke's law , Fick's law , and Ohm's law ) which serves to define the viscosity μ {\displaystyle \mu } . Its form is motivated by experiments which show that for a wide range of fluids, μ {\displaystyle \mu } is independent of strain rate. Such fluids are called Newtonian . Gases , water , and many common liquids can be considered Newtonian in ordinary conditions and contexts. However, there are many non-Newtonian fluids that significantly deviate from this behavior. For example: Trouton 's ratio

4641-494: Is particularly rich in carbohydrates, proteins and lipids. While their function is not always clear, and may be related to regulation of sugars, in most cases they appear to facilitate plant insect relationships. In contrast to floral nectaries, nectar produced outside the flower generally has a defensive function. The nectar attracts predatory insects which will eat both the nectar and any plant-eating insects around, thus functioning as "bodyguards". Foraging predatory insects show

4760-470: Is proportional to the fluid's viscosity. In general, viscosity depends on a fluid's state, such as its temperature, pressure, and rate of deformation. However, the dependence on some of these properties is negligible in certain cases. For example, the viscosity of a Newtonian fluid does not vary significantly with the rate of deformation. Zero viscosity (no resistance to shear stress ) is observed only at very low temperatures in superfluids ; otherwise,

4879-473: Is proportional to the percentage of fertilised ovules. For example, with watermelon , about a thousand grains of pollen must be delivered and spread evenly on the three lobes of the stigma to make a normal sized and shaped fruit. Outcrossing , or cross-fertilisation, and self-fertilisation represent different strategies with differing benefits and costs. An estimated 48.7% of plant species are either dioecious or self-incompatible obligate outcrossers. It

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4998-588: Is the dynamic viscosity of the fluid, often simply referred to as the viscosity . It is denoted by the Greek letter mu ( μ ). The dynamic viscosity has the dimensions ( m a s s / l e n g t h ) / t i m e {\displaystyle \mathrm {(mass/length)/time} } , therefore resulting in the SI units and the derived units : The aforementioned ratio u / y {\displaystyle u/y}

5117-408: Is the density, J {\displaystyle \mathbf {J} } and q {\displaystyle \mathbf {q} } are the mass and heat fluxes, and D {\displaystyle D} and k t {\displaystyle k_{t}} are the mass diffusivity and thermal conductivity. The fact that mass, momentum, and energy (heat) transport are among

5236-425: Is the local shear velocity. This expression is referred to as Newton's law of viscosity . In shearing flows with planar symmetry, it is what defines μ {\displaystyle \mu } . It is a special case of the general definition of viscosity (see below), which can be expressed in coordinate-free form. Use of the Greek letter mu ( μ {\displaystyle \mu } ) for

5355-413: Is the process in angiosperms (flowering plants) in which two sperm from each pollen tube fertilise two cells in a female gametophyte (sometimes called an embryo sac) that is inside an ovule. After the pollen tube enters the gametophyte, the pollen tube nucleus disintegrates and the two sperm cells are released; one of the two sperm cells fertilises the egg cell (at the bottom of the gametophyte near

5474-611: Is the ratio of extensional viscosity to shear viscosity . For a Newtonian fluid, the Trouton ratio is 3. Shear-thinning liquids are very commonly, but misleadingly, described as thixotropic. Viscosity may also depend on the fluid's physical state (temperature and pressure) and other, external , factors. For gases and other compressible fluids , it depends on temperature and varies very slowly with pressure. The viscosity of some fluids may depend on other factors. A magnetorheological fluid , for example, becomes thicker when subjected to

5593-554: Is the unit tensor. This equation can be thought of as a generalized form of Newton's law of viscosity. The bulk viscosity (also called volume viscosity) expresses a type of internal friction that resists the shearless compression or expansion of a fluid. Knowledge of κ {\displaystyle \kappa } is frequently not necessary in fluid dynamics problems. For example, an incompressible fluid satisfies ∇ ⋅ v = 0 {\displaystyle \nabla \cdot \mathbf {v} =0} and so

5712-456: Is to attract potential pollinators , which may include insects, including bees and moths , and vertebrates such as hummingbirds and bats . Nectaries can occur on any floral part, but they may also represent a modified part or a novel structure. The different types of floral nectaries include: Most members of Lamiaceae have a nectariferous disc which surrounds the ovary base and derived from developing ovarian tissue. In most Brassicaceae ,

5831-601: Is usually identical save for a fraction of their genes; each gamete is therefore genetically unique. At fertilisation, parental chromosomes combine. In humans , (2²²)² = 17.6x10 chromosomally different zygotes are possible for the non-sex chromosomes, even assuming no chromosomal crossover . If crossover occurs once, then on average (4²²)² = 309x10 genetically different zygotes are possible for every couple, not considering that crossover events can take place at most points along each chromosome. The X and Y chromosomes undergo no crossover events and are therefore excluded from

5950-1211: Is variable due to many factors, including flower age, plant location, and habitat management. Extrafloral nectaries (also known as extranuptial nectaries) are specialised nectar-secreting plant glands that develop outside of flowers and are not involved in pollination , generally on the leaf or petiole (foliar nectaries) and often in relation to the leaf venation . They are highly diverse in form, location, size, and mechanism. They have been described in virtually all above-ground plant parts—including stipules , cotyledons , fruits , and stems , among others. They range from single-celled trichomes to complex cup-like structures that may or may not be vascularized . Like floral nectaries, they consist of groups of glandular trichomes (e.g., Hibiscus spp.) or elongated secretory epidermal cells. The latter are often associated with underlying vascular tissue. They may be associated with specialised pockets ( domatia ), pits or raised regions (e.g., Euphorbiaceae ). The leaves of some tropical eudicots (e.g., Fabaceae ) and magnoliids (e.g., Piperaceae ) possess pearl glands or bodies which are globular trichomes specialised to attract ants. They secrete matter that

6069-559: The conceptus and such medical literature refers to the "products of conception" as the post-implantation embryo and its surrounding membranes. The term "conception" is not usually used in scientific literature because of its variable definition and connotation. Insects in different groups, including the Odonata ( dragonflies and damselflies ) and the Hymenoptera ( ants , bees , and wasps ) practise delayed fertilisation. Among

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6188-852: The Cenozoic , with weak support for a role played by arthropod herbivore diversifications. They are absent in bryophytes , gymnosperms , early angiosperms , magnoliids , and members of Apiales among the eudicots. Phylogenetic studies and the wide distribution of extrafloral nectaries among vascular plants point to multiple independent evolutionary origins of extrafloral nectaries in at least 457 independent lineages. The main ingredients in nectar are sugars in varying proportions of sucrose , glucose , and fructose . In addition, nectars have diverse other phytochemicals serving to both attract pollinators and discourage predators. Carbohydrates , amino acids , and volatiles function to attract some species, whereas alkaloids and polyphenols appear to provide

6307-466: The Latin viscum (" mistletoe "). Viscum also referred to a viscous glue derived from mistletoe berries. In materials science and engineering , there is often interest in understanding the forces or stresses involved in the deformation of a material. For instance, if the material were a simple spring, the answer would be given by Hooke's law , which says that the force experienced by

6426-589: The Zahn cup and the Ford viscosity cup —with the usage of each type varying mainly according to the industry. Also used in coatings, a Stormer viscometer employs load-based rotation to determine viscosity. The viscosity is reported in Krebs units (KU), which are unique to Stormer viscometers. Vibrating viscometers can also be used to measure viscosity. Resonant, or vibrational viscometers work by creating shear waves within

6545-425: The density of the fluid ( ρ ). It is usually denoted by the Greek letter nu ( ν ): and has the dimensions ( l e n g t h ) 2 / t i m e {\displaystyle \mathrm {(length)^{2}/time} } , therefore resulting in the SI units and the derived units : In very general terms, the viscous stresses in a fluid are defined as those resulting from

6664-436: The fruit . With multi-seeded fruits, multiple grains of pollen are necessary for syngamy with each ovule. The growth of the pollen tube is controlled by the vegetative (or tube) cytoplasm. Hydrolytic enzymes are secreted by the pollen tube that digest the female tissue as the tube grows down the stigma and style; the digested tissue is used as a nutrient source for the pollen tube as it grows. During pollen tube growth towards

6783-583: The gynoecium from certain pathogens. Floral nectaries have evolved and diverged into the different types of nectaries due to the various pollinators that visit the flowers. In Melastomataceae , different types of floral nectaries have evolved and been lost many times. Flowers that ancestrally produced nectar and had nectaries may have lost their ability to produce nectar due to a lack of nectar consumption by pollinators, such as certain species of bees . Instead they focused on energy allocation to pollen production. Species of angiosperms that have nectaries use

6902-405: The second law of thermodynamics requires all fluids to have positive viscosity. A fluid that has zero viscosity (non-viscous) is called ideal or inviscid . For non-Newtonian fluid 's viscosity, there are pseudoplastic , plastic , and dilatant flows that are time-independent, and there are thixotropic and rheopectic flows that are time-dependent. The word "viscosity" is derived from

7021-431: The shear viscosity . However, at least one author discourages the use of this terminology, noting that μ {\displaystyle \mu } can appear in non-shearing flows in addition to shearing flows. In fluid dynamics, it is sometimes more appropriate to work in terms of kinematic viscosity (sometimes also called the momentum diffusivity ), defined as the ratio of the dynamic viscosity ( μ ) over

7140-491: The BG and EE systems. Nonstandard units include the reyn (lbf·s/in ), a British unit of dynamic viscosity. In the automotive industry the viscosity index is used to describe the change of viscosity with temperature. The reciprocal of viscosity is fluidity , usually symbolized by ϕ = 1 / μ {\displaystyle \phi =1/\mu } or F = 1 / μ {\displaystyle F=1/\mu } , depending on

7259-468: The Couette flow, a fluid is trapped between two infinitely large plates, one fixed and one in parallel motion at constant speed u {\displaystyle u} (see illustration to the right). If the speed of the top plate is low enough (to avoid turbulence), then in steady state the fluid particles move parallel to it, and their speed varies from 0 {\displaystyle 0} at

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7378-506: The Odonata, females may mate with multiple males, and store sperm until the eggs are laid. The male may hover above the female during egg-laying (oviposition) to prevent her from mating with other males and replacing his sperm; in some groups such as the darters, the male continues to grasp the female with his claspers during egg-laying, the pair flying around in tandem. Among social Hymenoptera, honeybee queens mate only on mating flights, in

7497-469: The Vegetable Kingdom (pages 466-467) summed up his findings in the following way. "It has been shown in the present volume that the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigour and fertility over the self-fertilised offspring from one of

7616-557: The ZP3 and is important for binding with the sperm and activating the acrosome reaction. ZP3 is sufficient though unnecessary for sperm/egg binding. Two additional sperm receptors exist: a 250kD protein that binds to an oviduct secreted protein, and SED1, which independently binds to the zona. After the acrosome reaction, the sperm is believed to remain bound to the zona pellucida through exposed ZP2 receptors. These receptors are unknown in mice but have been identified in guinea pigs. In mammals,

7735-469: The ability to store sperm for extended periods of time and can fertilise their eggs at their own desire. Oviparous animals producing eggs with thin tertiary membranes or no membranes at all, on the other hand, use external fertilisation methods. Such animals may be more precisely termed ovuliparous. External fertilisation is advantageous in that it minimises contact (which decreases the risk of disease transmission), and greater genetic variation. Sperm find

7854-406: The actual persistence of meiosis and self-fertilisation as a form of reproduction in long-established self-fertilising plants may be related to the immediate benefit of efficient recombinational repair of DNA damage during formation of germ cells provided by meiosis at each generation. The mechanics behind fertilisation has been studied extensively in sea urchins and mice. This research addresses

7973-400: The advantage of reproductive assurance . Self-fertilisation can therefore result in improved colonisation ability. In some species, self-fertilisation has persisted over many generations. Capsella rubella is a self-fertilising species that became self-compatible 50,000 to 100,000 years ago. Arabidopsis thaliana is a predominantly self-fertilising plant with an out-crossing rate in

8092-531: The arithmetic and the reference table provided in ASTM D 2161. Fertilization This is an accepted version of this page Fertilisation or fertilization (see spelling differences ), also known as generative fertilisation , syngamy and impregnation , is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or offspring. While processes such as insemination or pollination , which happen before

8211-467: The binding of the spermatozoon to the GalT initiates the acrosome reaction . This process releases the hyaluronidase that digests the matrix of hyaluronic acid in the vestments around the oocyte. Additionally, heparin-like glycosaminoglycans (GAGs) are released near the oocyte that promote the acrosome reaction. Fusion between the oocyte plasma membranes and sperm follows and allows the sperm nucleus ,

8330-425: The bottom to u {\displaystyle u} at the top. Each layer of fluid moves faster than the one just below it, and friction between them gives rise to a force resisting their relative motion. In particular, the fluid applies on the top plate a force in the direction opposite to its motion, and an equal but opposite force on the bottom plate. An external force is therefore required in order to keep

8449-467: The calculation. The mitochondrial DNA is only inherited from the maternal parent. Shortly after the sperm fuse with the egg, the two sperm centrioles form the embryo first centrosome and microtubule aster . The sperm centriole, found near the male pronucleus, recruit egg Pericentriolar material proteins forming the zygote first centrosome. This centrosome nucleates microtubules in the shape of stars called astral microtubules. The microtubules span

8568-602: The convention used, measured in reciprocal poise (P , or cm · s · g ), sometimes called the rhe . Fluidity is seldom used in engineering practice. At one time the petroleum industry relied on measuring kinematic viscosity by means of the Saybolt viscometer , and expressing kinematic viscosity in units of Saybolt universal seconds (SUS). Other abbreviations such as SSU ( Saybolt seconds universal ) or SUV ( Saybolt universal viscosity ) are sometimes used. Kinematic viscosity in centistokes can be converted from SUS according to

8687-465: The dynamic viscosity (sometimes also called the absolute viscosity ) is common among mechanical and chemical engineers , as well as mathematicians and physicists. However, the Greek letter eta ( η {\displaystyle \eta } ) is also used by chemists, physicists, and the IUPAC . The viscosity μ {\displaystyle \mu } is sometimes also called

8806-555: The egg to develop without fertilisation or syngamy. The sperm may enter the egg. Hybridogenesis : One genome is eliminated to produce haploid eggs. Canina meiosis : (sometimes called "permanent odd polyploidy") one genome is transmitted in the Mendelian fashion, others are transmitted clonally. The major benefit of cross-fertilisation is generally thought to be the avoidance of inbreeding depression . Charles Darwin , in his 1876 book The Effects of Cross and Self Fertilisation in

8925-405: The eggs via chemotaxis , a type of ligand/receptor interaction. Resact is a 14 amino acid peptide purified from the jelly coat of A. punctulata that attracts the migration of sperm. After finding the egg, the sperm penetrates the jelly coat through a process called sperm activation. In another ligand/receptor interaction, an oligosaccharide component of the egg binds and activates a receptor on

9044-598: The elongated tissue and more easily reach the nectaries and obtain the nectar reward. Different characteristics of the spur, such as its length or position in the flower, may determine the type of pollinator that visits the flower. Defense from herbivory is often one of the roles of extrafloral nectaries. Floral nectaries can also be involved in defense. In addition to the sugars found in nectar, certain proteins may also be found in nectar secreted by floral nectaries. In tobacco plants, these proteins have antimicrobial and antifungal properties and can be secreted to defend

9163-407: The equivalent forms pascal - second (Pa·s), kilogram per meter per second (kg·m ·s ) and poiseuille (Pl). The CGS unit is the poise (P, or g·cm ·s = 0.1 Pa·s), named after Jean Léonard Marie Poiseuille . It is commonly expressed, particularly in ASTM standards, as centipoise (cP). The centipoise is convenient because the viscosity of water at 20 °C is about 1 cP, and one centipoise

9282-409: The fluid do not depend on the distance the fluid has been sheared; rather, they depend on how quickly the shearing occurs. Viscosity is the material property which relates the viscous stresses in a material to the rate of change of a deformation (the strain rate). Although it applies to general flows, it is easy to visualize and define in a simple shearing flow, such as a planar Couette flow . In

9401-459: The fluid is essential to obtain accurate measurements, particularly in materials like lubricants, whose viscosity can double with a change of only 5 °C. A rheometer is used for fluids that cannot be defined by a single value of viscosity and therefore require more parameters to be set and measured than is the case for a viscometer. For some fluids, the viscosity is constant over a wide range of shear rates ( Newtonian fluids ). The fluids without

9520-834: The forager after only a sip, motivating it to visit other plants, therefore maximizing the pollination efficiency gained by the plant for a minimum nectar output. Neurotoxins such as aesculin are present in some nectars such as that of the California buckeye . Nectar contains water, essential oils , carbohydrates , amino acids , ions , and numerous other compounds. Some insect pollinated plants lack nectaries, but attract pollinators through other secretory structures. Elaiophores are similar to nectaries but are oil secreting. Osmophores are modified structural structures that produce volatile scents. In orchids , these have pheromone qualities. Osmophores have thick domed or papillate epidermis and dense cytoplasm. Platanthera bifolia produces

9639-441: The formation of a diploid cell called a zygote . The zygote divides to form a blastocyst and, upon entering the uterus, implants in the endometrium, beginning pregnancy . Embryonic implantation not in the uterine wall results in an ectopic pregnancy that can kill the mother. In such animals as rabbits, coitus induces ovulation by stimulating the release of the pituitary hormone gonadotropin; this release greatly increases

9758-399: The formation of a fertilisation cone . Mammals internally fertilise through copulation . After a male ejaculates , many sperm move to the upper vagina (via contractions from the vagina) through the cervix and across the length of the uterus to meet the ovum. In cases where fertilisation occurs, the female usually ovulates during a period that extends from hours before copulation to

9877-479: The formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in a mode called by him as epigenetic . In 1784, Spallanzani established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs. In 1827, Karl Ernst von Baer observed a therian mammalian egg for the first time. Oscar Hertwig (1876), in Germany, described

9996-466: The fusion of gametes, are also sometimes informally referred to as fertilisation, these are technically separate processes. The cycle of fertilisation and development of new individuals is called sexual reproduction . During double fertilisation in angiosperms , the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation. In antiquity, Aristotle conceived

10115-432: The fusion of nuclei of spermatozoa and of ova from sea urchin . The evolution of fertilisation is related to the origin of meiosis , as both are part of sexual reproduction , originated in eukaryotes . One hypothesis states that meiosis originated from mitosis. The gametes that participate in fertilisation of plants are the sperm (male) and the egg (female) cell. Various plant groups have differing methods by which

10234-406: The gametes produced by the male and female gametophytes come together and are fertilised. In bryophytes and pteridophytic land plants, fertilisation of the sperm and egg takes place within the archegonium . In seed plants , the male gametophyte is formed within a pollen grain. After pollination , the pollen grain germinates , and a pollen tube grows and penetrates the ovule through

10353-893: The known occurrences. The families showing the most recorded occurrences of extrafloral nectaries are Fabaceae , with 1069 species, Passifloraceae , with 438 species, and Malvaceae , with 301 species. The genera with the most recorded occurrences are Passiflora (322 species, Passifloraceae), Inga (294 species, Fabaceae), and Acacia (204 species, Fabaceae). Other genera with extrafloral nectaries include Salix ( Salicaceae ), Prunus ( Rosaceae ) and Gossypium ( Malvaceae ). Foliar nectaries have also been observed in 101 species of ferns belonging to eleven genera and six families, most of them belonging to Cyatheales (tree ferns) and Polypodiales . Fern nectaries appear to have evolved around 135 million years ago, nearly simultaneously with angiosperms. However, fern nectaries did not diversify remarkably until nearly 100 million years later, in

10472-541: The likelihood of pregnancy. Fertilisation in humans is the union of a human egg and sperm , usually occurring in the ampulla of the fallopian tube , producing a single celled zygote , the first stage of life in the development of a genetically unique organism, and initiating embryonic development . Scientists discovered the dynamics of human fertilisation in the nineteenth century. The term conception commonly refers to "the process of becoming pregnant involving fertilisation or implantation or both". Its use makes it

10591-529: The liquid. In this method, the sensor is submerged in the fluid and is made to resonate at a specific frequency. As the surface of the sensor shears through the liquid, energy is lost due to its viscosity. This dissipated energy is then measured and converted into a viscosity reading. A higher viscosity causes a greater loss of energy. Extensional viscosity can be measured with various rheometers that apply extensional stress . Volume viscosity can be measured with an acoustic rheometer . Apparent viscosity

10710-411: The micropyle), forming a diploid (2n) zygote . This is the point when fertilisation actually occurs; pollination and fertilisation are two separate processes. The nucleus of the other sperm cell fuses with two haploid polar nuclei (contained in the central cell) in the centre of the gametophyte. The resulting cell is triploid (3n). This triploid cell divides through mitosis and forms the endosperm ,

10829-435: The most relevant processes in continuum mechanics is not a coincidence: these are among the few physical quantities that are conserved at the microscopic level in interparticle collisions. Thus, rather than being dictated by the fast and complex microscopic interaction timescale, their dynamics occurs on macroscopic timescales, as described by the various equations of transport theory and hydrodynamics. Newton's law of viscosity

10948-415: The mother, or in some cases genetically differ from her but inherit only part of her DNA. Parthenogenesis occurs in many plants and animals and may be induced in others through a chemical or electrical stimulus to the egg cell. In 2004, Japanese researchers led by Tomohiro Kono succeeded after 457 attempts to merge the ova of two mice by blocking certain proteins that would normally prevent the possibility;

11067-452: The nectar to attract pollinators that consume the nectar, such as birds and butterflies . In Bromeliaceae , septal nectaries (a form of gynoecial nectary) are common in species that are insect or bird pollinated. In species that are wind pollinated, nectaries are often absent because there is no pollinator. In flowers that are generally pollinated by a long-tongued organism such as certain flies , moths , butterflies, and birds, nectaries in

11186-418: The nectary is at the base of the stamen filament. Many monocotyledons have septal nectaries, which are at the unfused margins of the carpels. These exude nectar from small pores on the surface of the gynoecium. Nectaries may also vary in color, number, and symmetry. Nectaries can also be categorized as structural or non-structural. Structural nectaries refer to specific areas of tissue that exude nectar, such as

11305-426: The ovaries are common because they are able to reach the nectar reward when pollinating. Sepal and petal nectaries are often more common in species that are pollinated by short-tongued insects that cannot reach so far into the flower. Nectar secretion increases as the flower is visited by pollinators. After pollination, the nectar is frequently reabsorbed into the plant. The amount of nectar in flowers at any given time

11424-421: The ovary, the generative nucleus divides to produce two separate sperm nuclei (haploid number of chromosomes) – a growing pollen tube therefore contains three separate nuclei, two sperm and one tube. The sperms are interconnected and dimorphic, the large one, in a number of plants, is also linked to the tube nucleus and the interconnected sperm and the tube nucleus form the "male germ unit". Double fertilisation

11543-444: The oviduct and the ampulla, and chemotactic gradients of progesterone have been confirmed as the signal emanating from the cumulus oophorus cells surrounding rabbit and human oocytes. Capacitated and hyperactivated sperm respond to these gradients by changing their behaviour and moving towards the cumulus-oocyte complex. Other chemotactic signals such as formyl Met-Leu-Phe (fMLF) may also guide spermatozoa. The zona pellucida ,

11662-452: The plant as they brush against the reproductive organs, the stamen and pistil , of the plant and pick up or deposit pollen . Nectar from floral nectaries is sometimes used as a reward to insects, such as ants , that protect the plant from predators. Many floral families have evolved a nectar spur . These spurs are projections of various lengths formed from different tissues, such as the petals or sepals. They allow for pollinators to land on

11781-449: The pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed a family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium,

11900-491: The pollen tube, causing these channels to take up Calcium ions in large amounts. This increased uptake of calcium causes the pollen tube to rupture, and release its sperm into the ovule. Pistil feeding assays in which plants were fed diphenyl iodonium chloride (DPI) suppressed ROS concentrations in Arabidopsis , which in turn prevented pollen tube rupture. After being fertilised, the ovary starts to swell and develop into

12019-546: The question of how the sperm and the appropriate egg find each other and the question of how only one sperm gets into the egg and delivers its contents. There are three steps to fertilisation that ensure species-specificity: Consideration as to whether an animal (more specifically a vertebrate) uses internal or external fertilisation is often dependent on the method of birth. Oviparous animals laying eggs with thick calcium shells, such as chickens , or thick leathery shells generally reproduce via internal fertilisation so that

12138-515: The relative velocity of different fluid particles. As such, the viscous stresses must depend on spatial gradients of the flow velocity. If the velocity gradients are small, then to a first approximation the viscous stresses depend only on the first derivatives of the velocity. (For Newtonian fluids, this is also a linear dependence.) In Cartesian coordinates, the general relationship can then be written as where μ i j k ℓ {\displaystyle \mu _{ijk\ell }}

12257-465: The resulting embryo normally developed into a mouse. Allogamy , which is also known as cross-fertilisation, refers to the fertilisation of an egg cell from one individual with the male gamete of another. Autogamy which is also known as self-fertilisation, occurs in such hermaphroditic organisms as plants and flatworms; therein, two gametes from one individual fuse. Some relatively unusual forms of reproduction are: Gynogenesis : A sperm stimulates

12376-445: The secretory region, where it is secreted from the cells through vesicles packaged by the endoplasmic reticulum . The adjacent subepidermal cells may also be secretory. Flowers that have longer nectaries sometimes have a vascular strand in the nectary to assist in transport over a longer distance. Pollinators feed on the nectar and depending on the location of the nectary the pollinator assists in fertilization and outcrossing of

12495-435: The sperm and causes the acrosomal reaction . The acrosomal vesicles of the sperm fuse with the plasma membrane and are released. In this process, molecules bound to the acrosomal vesicle membrane, such as bindin, are exposed on the surface of the sperm. These contents digest the jelly coat and eventually the vitelline membrane. In addition to the release of acrosomal vesicles, there is explosive polymerisation of actin to form

12614-501: The sperm fertilises the egg without having to pass through the thick, protective, tertiary layer of the egg. Ovoviviparous and viviparous animals also use internal fertilisation. Although some organisms reproduce via amplexus , they may still use internal fertilisation, as with some salamanders. Advantages of internal fertilisation include minimal waste of gametes, greater chance of individual egg fertilisation, longer period of egg protection, and selective fertilisation. Many females have

12733-555: The term fugitive elasticity for fluid viscosity. However, many liquids (including water) will briefly react like elastic solids when subjected to sudden stress. Conversely, many "solids" (even granite ) will flow like liquids, albeit very slowly, even under arbitrarily small stress. Such materials are best described as viscoelastic —that is, possessing both elasticity (reaction to deformation) and viscosity (reaction to rate of deformation). Viscoelastic solids may exhibit both shear viscosity and bulk viscosity. The extensional viscosity

12852-715: The term containing κ {\displaystyle \kappa } drops out. Moreover, κ {\displaystyle \kappa } is often assumed to be negligible for gases since it is 0 {\displaystyle 0} in a monatomic ideal gas . One situation in which κ {\displaystyle \kappa } can be important is the calculation of energy loss in sound and shock waves , described by Stokes' law of sound attenuation , since these phenomena involve rapid expansions and compressions. The defining equations for viscosity are not fundamental laws of nature, so their usefulness, as well as methods for measuring or calculating

12971-583: The top plate moving at constant speed. In many fluids, the flow velocity is observed to vary linearly from zero at the bottom to u {\displaystyle u} at the top. Moreover, the magnitude of the force, F {\displaystyle F} , acting on the top plate is found to be proportional to the speed u {\displaystyle u} and the area A {\displaystyle A} of each plate, and inversely proportional to their separation y {\displaystyle y} : The proportionality factor

13090-495: The tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vivo pollinated pistils, and pollen tubes were observed to immediately extend toward the proteins. Transgenic plants lacking the ability to produce TTS proteins had slower pollen tube growth and reduced fertility. The rupture of the pollen tube to release sperm in Arabidopsis has been shown to depend on

13209-513: The two systems differ only in how force and mass are defined. In the BG system the pound is a basic unit from which the unit of mass (the slug ) is defined by Newton's Second Law , whereas in the EE system the units of force and mass (the pound-force and pound-mass respectively) are defined independently through the Second Law using the proportionality constant g c . Kinematic viscosity has units of square feet per second (ft /s) in both

13328-451: The types of floral nectaries previously listed. Non-structural nectaries secrete nectar infrequently from non-differentiated tissues. The different types of floral nectaries coevolved depending on the pollinator that feeds on the plant's nectar. Nectar is secreted from epidermal cells of the nectaries, which have a dense cytoplasm , by means of trichomes or modified stomata . Adjacent vascular tissue conducts phloem bringing sugars to

13447-488: The typical centriole , and atypical centriole that is attached to the flagellum , but not the mitochondria , to enter the oocyte. The protein CD9 likely mediates this fusion in mice (the binding homolog). The egg " activates " itself upon fusing with a single sperm cell and thereby changes its cell membrane to prevent fusion with other sperm. Zinc atoms are released during this activation. This process ultimately leads to

13566-403: The viscosity depends only space- and time-dependent macroscopic fields (such as temperature and density) defining local equilibrium. Nevertheless, viscosity may still carry a non-negligible dependence on several system properties, such as temperature, pressure, and the amplitude and frequency of any external forcing. Therefore, precision measurements of viscosity are only defined with respect to

13685-571: The viscosity rank-2 tensor is isotropic reduces these 81 coefficients to three independent parameters α {\displaystyle \alpha } , β {\displaystyle \beta } , γ {\displaystyle \gamma } : and furthermore, it is assumed that no viscous forces may arise when the fluid is undergoing simple rigid-body rotation, thus β = γ {\displaystyle \beta =\gamma } , leaving only two independent parameters. The most usual decomposition

13804-500: The viscosity, must be established using separate means. A potential issue is that viscosity depends, in principle, on the full microscopic state of the fluid, which encompasses the positions and momenta of every particle in the system. Such highly detailed information is typically not available in realistic systems. However, under certain conditions most of this information can be shown to be negligible. In particular, for Newtonian fluids near equilibrium and far from boundaries (bulk state),

13923-534: The whole valium of the egg, allowing the egg pronucleus to use the cables to get to the male pronucleus. As the male and female pronuclei approach each other, the single centrosome split into two centrosomes located in the interphase between the pronuclei. Then the centrosome via the astral microtubules polarises the genome inside the pronuclei. Organisms that normally reproduce sexually can also reproduce via parthenogenesis , wherein an unfertilised female gamete produces viable offspring. These offspring may be clones of

14042-537: The wild of less than 0.3%; a study suggested that self-fertilisation evolved roughly a million years ago or more in A. thaliana . In long-established self-fertilising plants, the masking of deleterious mutations and the production of genetic variability is infrequent and thus unlikely to provide a sufficient benefit over many generations to maintain the meiotic apparatus. Consequently, one might expect self-fertilisation to be replaced in nature by an ameiotic asexual form of reproduction that would be less costly. However

14161-513: The word from νε- or νη- "not" plus κτα- or κτεν- "kill", meaning "unkillable", thus "immortal". The common use of the word "nectar" to refer to the "sweet liquid in flowers", is first recorded in AD 1600. A nectary or honey gland is floral tissue found in different locations in the flower and is one of several secretory floral structures, including elaiophores and osmophores, producing nectar, oil and scent respectively. The function of these structures

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