Photosystems are functional and structural units of protein complexes involved in photosynthesis . Together they carry out the primary photochemistry of photosynthesis : the absorption of light and the transfer of energy and electrons . Photosystems are found in the thylakoid membranes of plants, algae, and cyanobacteria. These membranes are located inside the chloroplasts of plants and algae, and in the cytoplasmic membrane of photosynthetic bacteria. There are two kinds of photosystems: PSI and PSII.
68-473: PSII will absorb red light, and PSI will absorb far-red light. Although photosynthetic activity will be detected when the photosystems are exposed to either red or far-red light, the photosynthetic activity will be the greatest when plants are exposed to both wavelengths of light. Studies have actually demonstrated that the two wavelengths together have a synergistic effect on the photosynthetic activity, rather than an additive one. Each photosystem has two parts:
136-418: A can also be found in very small quantities in the green sulfur bacteria , an anaerobic photoautotroph . These organisms use bacteriochlorophyll and some chlorophyll a but do not produce oxygen. Anoxygenic photosynthesis is the term applied to this process, unlike oxygenic photosynthesis where oxygen is produced during the light reactions of photosynthesis . The molecular structure of chlorophyll
204-421: A consists of a chlorin ring, whose four nitrogen atoms surround a central magnesium atom, and has several other attached side chains and a hydrocarbon tail formed by a phytol ester . Chlorophyll a contains a magnesium ion encased in a large ring structure known as a chlorin . The chlorin ring is a heterocyclic compound derived from pyrrole . Four nitrogen atoms from the chlorin surround and bind
272-411: A local wavelength . An example is shown in the figure. In general, the envelope of the wave packet moves at a speed different from the constituent waves. Using Fourier analysis , wave packets can be analyzed into infinite sums (or integrals) of sinusoidal waves of different wavenumbers or wavelengths. Louis de Broglie postulated that all particles with a specific value of momentum p have
340-582: A quinone terminal electron acceptor. Both reaction center types are present in chloroplasts and cyanobacteria, and work together to form a unique photosynthetic chain able to extract electrons from water, creating oxygen as a byproduct. A reaction center comprises several (about 25-30) protein subunits, which provide a scaffold for a series of cofactors. The cofactors can be pigments (like chlorophyll , pheophytin , carotenoids ), quinones, or iron-sulfur clusters . Each photosystem has two main subunits: an antenna complex (a light harvesting complex or LHC) and
408-407: A wave or periodic function is the distance over which the wave's shape repeats. In other words, it is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, troughs, or zero crossings . Wavelength is a characteristic of both traveling waves and standing waves , as well as other spatial wave patterns. The inverse of the wavelength
476-455: A circular aperture, the diffraction-limited image spot is known as an Airy disk ; the distance x in the single-slit diffraction formula is replaced by radial distance r and the sine is replaced by 2 J 1 , where J 1 is a first order Bessel function . The resolvable spatial size of objects viewed through a microscope is limited according to the Rayleigh criterion , the radius to
544-490: A common ancestor, but have since diversified. Each of the photosystem can be identified by the wavelength of light to which it is most reactive (700 nanometers for PSI and 680 nanometers for PSII in chloroplasts), the amount and type of light-harvesting complex present, and the type of terminal electron acceptor used. Type I photosystems use ferredoxin -like iron-sulfur cluster proteins as terminal electron acceptors, while type II photosystems ultimately shuttle electrons to
612-468: A linear system the sinusoid is the unique shape that propagates with no shape change – just a phase change and potentially an amplitude change. The wavelength (or alternatively wavenumber or wave vector ) is a characterization of the wave in space, that is functionally related to its frequency, as constrained by the physics of the system. Sinusoids are the simplest traveling wave solutions, and more complex solutions can be built up by superposition . In
680-461: A reaction center, where the photochemistry occurs, and an antenna complex , which surrounds the reaction center. The antenna complex contains hundreds of chlorophyll molecules which funnel the excitation energy to the center of the photosystem. At the reaction center, the energy will be trapped and transferred to produce a high energy molecule. The main function of PSII is to efficiently split water into oxygen molecules and protons. PSII will provide
748-473: A reaction center. The antenna complex is where light is captured, while the reaction center is where this light energy is transformed into chemical energy. At the reaction center, there are many polypeptides that are surrounded by pigment proteins. At the center of the reaction center is a special pair of chlorophyll molecules. Each PSII has about 8 LHCII. These contain about 14 chlorophyll a and chlorophyll b molecules, as well as about four carotenoids . In
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#1732773357745816-426: A regular lattice. This produces aliasing because the same vibration can be considered to have a variety of different wavelengths, as shown in the figure. Descriptions using more than one of these wavelengths are redundant; it is conventional to choose the longest wavelength that fits the phenomenon. The range of wavelengths sufficient to provide a description of all possible waves in a crystalline medium corresponds to
884-432: A result, the change in direction upon entering a different medium changes with the wavelength of the wave. For electromagnetic waves the speed in a medium is governed by its refractive index according to where c is the speed of light in vacuum and n ( λ 0 ) is the refractive index of the medium at wavelength λ 0 , where the latter is measured in vacuum rather than in the medium. The corresponding wavelength in
952-491: A steady stream of electrons to PSI, which will boost these in energy and transfer them to NADP and H to make NADPH . The hydrogen from this NADPH can then be used in a number of different processes within the plant. Reaction centers are multi-protein complexes found within the thylakoid membrane. At the heart of a photosystem lies the reaction center , which is an enzyme that uses light to reduce and oxidize molecules (give off and take up electrons). This reaction center
1020-404: A traveling wave. For example, the speed of light can be determined from observation of standing waves in a metal box containing an ideal vacuum. Traveling sinusoidal waves are often represented mathematically in terms of their velocity v (in the x direction), frequency f and wavelength λ as: where y is the value of the wave at any position x and time t , and A is the amplitude of
1088-531: A wavelength λ = h / p , where h is the Planck constant . This hypothesis was at the basis of quantum mechanics . Nowadays, this wavelength is called the de Broglie wavelength . For example, the electrons in a CRT display have a De Broglie wavelength of about 10 m . To prevent the wave function for such a particle being spread over all space, de Broglie proposed using wave packets to represent particles that are localized in space. The spatial spread of
1156-424: Is also responsible for the familiar phenomenon in which light is separated into component colours by a prism . Separation occurs when the refractive index inside the prism varies with wavelength, so different wavelengths propagate at different speeds inside the prism, causing them to refract at different angles. The mathematical relationship that describes how the speed of light within a medium varies with wavelength
1224-464: Is an undulatory motion that stays in one place. A sinusoidal standing wave includes stationary points of no motion, called nodes , and the wavelength is twice the distance between nodes. The upper figure shows three standing waves in a box. The walls of the box are considered to require the wave to have nodes at the walls of the box (an example of boundary conditions ), thus determining the allowed wavelengths. For example, for an electromagnetic wave, if
1292-475: Is approximately 500mV, while the E m for P680 is approximately 1,100-1,200 mV. Chlorophyll a is very important in the energy phase of photosynthesis. Two electrons need to be passed to an electron acceptor for the process of photosynthesis to proceed. Within the reaction centers of both photosystems there are a pair of chlorophyll a molecules that pass electrons on to the transport chain through redox reactions. The concentration of chlorophyll A
1360-403: Is called diffraction . Two types of diffraction are distinguished, depending upon the separation between the source and the screen: Fraunhofer diffraction or far-field diffraction at large separations and Fresnel diffraction or near-field diffraction at close separations. In the analysis of the single slit, the non-zero width of the slit is taken into account, and each point in the aperture
1428-459: Is called the spatial frequency . Wavelength is commonly designated by the Greek letter lambda ( λ ). The term "wavelength" is also sometimes applied to modulated waves, and to the sinusoidal envelopes of modulated waves or waves formed by interference of several sinusoids. Assuming a sinusoidal wave moving at a fixed wave speed, wavelength is inversely proportional to the frequency of
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#17327733577451496-495: Is described by the Jacobi elliptic function of m th order, usually denoted as cn ( x ; m ) . Large-amplitude ocean waves with certain shapes can propagate unchanged, because of properties of the nonlinear surface-wave medium. If a traveling wave has a fixed shape that repeats in space or in time, it is a periodic wave . Such waves are sometimes regarded as having a wavelength even though they are not sinusoidal. As shown in
1564-413: Is known as a dispersion relation . Wavelength can be a useful concept even if the wave is not periodic in space. For example, in an ocean wave approaching shore, shown in the figure, the incoming wave undulates with a varying local wavelength that depends in part on the depth of the sea floor compared to the wave height. The analysis of the wave can be based upon comparison of the local wavelength with
1632-427: Is large compared to the slit separation d ) then the paths are nearly parallel, and the path difference is simply d sin θ . Accordingly, the condition for constructive interference is: where m is an integer, and for destructive interference is: Thus, if the wavelength of the light is known, the slit separation can be determined from the interference pattern or fringes , and vice versa . For multiple slits,
1700-488: Is related to position x via a squared sinc function : where L is the slit width, R is the distance of the pattern (on the screen) from the slit, and λ is the wavelength of light used. The function S has zeros where u is a non-zero integer, where are at x values at a separation proportion to wavelength. Diffraction is the fundamental limitation on the resolving power of optical instruments, such as telescopes (including radiotelescopes ) and microscopes . For
1768-434: Is surrounded by light-harvesting complexes that enhance the absorption of light. In addition, surrounding the reaction center are pigments which will absorb light. The pigments which absorb light at the highest energy level are found furthest from the reaction center. On the other hand, the pigments with the lowest energy level are more closely associated with the reaction center. Energy will be efficiently transferred from
1836-458: Is taken as the source of one contribution to the beam of light ( Huygens' wavelets ). On the screen, the light arriving from each position within the slit has a different path length, albeit possibly a very small difference. Consequently, interference occurs. In the Fraunhofer diffraction pattern sufficiently far from a single slit, within a small-angle approximation , the intensity spread S
1904-401: Is the enzyme that completes the biosynthesis of chlorophyll a by catalysing the reaction EC 2.5.1.62 This forms an ester of the carboxylic acid group in chlorophyllide a with the 20-carbon diterpene alcohol phytol . Chlorophyll a absorbs light within the violet , blue and red wavelengths. Accessory photosynthetic pigments broaden the spectrum of light absorbed, increasing
1972-417: Is used in the interferometer . A simple example is an experiment due to Young where light is passed through two slits . As shown in the figure, light is passed through two slits and shines on a screen. The path of the light to a position on the screen is different for the two slits, and depends upon the angle θ the path makes with the screen. If we suppose the screen is far enough from the slits (that is, s
2040-406: The antenna complex , ending in the reaction center where specific chlorophylls P680 and P700 are located. Chlorophyll a is essential for most photosynthetic organisms to release chemical energy but is not the only pigment that can be used for photosynthesis. All oxygenic photosynthetic organisms use chlorophyll a , but differ in accessory pigments like chlorophyll b . Chlorophyll
2108-411: The chloroplast strikes the pigments in the thylakoid membrane and excites their electrons. Since the chlorophyll a molecules only capture certain wavelengths, organisms may use accessory pigments to capture a wider range of light energy shown as the yellow circles. It then transfers captured light from one pigment to the next as resonance energy, passing energy one pigment to the other until reaching
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2176-422: The cosine phase instead of the sine phase when describing a wave is based on the fact that the cosine is the real part of the complex exponential in the wave The speed of a wave depends upon the medium in which it propagates. In particular, the speed of light in a medium is less than in vacuum , which means that the same frequency will correspond to a shorter wavelength in the medium than in vacuum, as shown in
2244-443: The speed of sound is 343 m/s (at room temperature and atmospheric pressure ). The wavelengths of sound frequencies audible to the human ear (20 Hz –20 kHz) are thus between approximately 17 m and 17 mm , respectively. Somewhat higher frequencies are used by bats so they can resolve targets smaller than 17 mm. Wavelengths in audible sound are much longer than those in visible light. A standing wave
2312-444: The thylakoid membrane . Energy from PSI drives this process and is harnessed (the whole process is termed chemiosmosis ) to pump protons across the membrane, into the thylakoid lumen space from the chloroplast stroma. This will provide a potential energy difference between lumen and stroma, which amounts to a proton-motive force that can be utilized by the proton-driven ATP synthase to generate ATP. If electrons only pass through once,
2380-466: The absorption spectrum of light. For instance, the only difference between chlorophyll a and chlorophyll b is that chlorophyll b has an aldehyde instead of a methyl group at the C-7 position. The phytol ester of chlorophyll a (R in the diagram) is a long hydrophobic tail which anchors the molecule to other hydrophobic proteins in the thylakoid membrane of the chloroplast . Once detached from
2448-399: The box has ideal conductive walls, the condition for nodes at the walls results because the conductive walls cannot support a tangential electric field, forcing the wave to have zero amplitude at the wall. The stationary wave can be viewed as the sum of two traveling sinusoidal waves of oppositely directed velocities. Consequently, wavelength, period, and wave velocity are related just as for
2516-485: The central bright portion (radius to first null of the Airy disk ) of the image diffracted by a circular aperture, a measure most commonly used for telescopes and cameras, is: Chlorophyll a Chlorophyll a is a specific form of chlorophyll used in oxygenic photosynthesis . It absorbs most energy from wavelengths of violet-blue and orange-red light, and it is a poor absorber of green and near-green portions of
2584-527: The chlorophyll and boosts its energy further, enough that it can split water in the oxygen evolving complex (OEC) of PSII and recover its electron. At the heart of the OEC are 4 Mn atoms, each of which can trap one electron. The electrons harvested from the splitting of two waters fill the OEC complex in its highest-energy state, which holds 4 excess electrons. Electrons travel through the cytochrome b6f complex to photosystem I via an electron transport chain within
2652-569: The direction and wavenumber of a plane wave in 3-space , parameterized by position vector r . In that case, the wavenumber k , the magnitude of k , is still in the same relationship with wavelength as shown above, with v being interpreted as scalar speed in the direction of the wave vector. The first form, using reciprocal wavelength in the phase, does not generalize as easily to a wave in an arbitrary direction. Generalizations to sinusoids of other phases, and to complex exponentials, are also common; see plane wave . The typical convention of using
2720-671: The electrons were not transferred away after excitation to a high energy state, they would lose energy by fluorescence back to the ground state, which would not allow plants to drive photosynthesis.) The reaction center will drive photosynthesis by taking light and turning it into chemical energy that can then be used by the chloroplast. Two families of reaction centers in photosystems can be distinguished: type I reaction centers (such as photosystem I ( P700 ) in chloroplasts and in green-sulfur bacteria) and type II reaction centers (such as photosystem II ( P680 ) in chloroplasts and in non-sulfur purple bacteria). The two photosystems originated from
2788-522: The enzyme NADP reductase. Electrons and protons are added to NADP to form NADPH. This reducing (hydrogenation) agent is transported to the Calvin cycle to react with glycerate 3-phosphate , along with ATP to form glyceraldehyde 3-phosphate , the basic building block from which plants can make a variety of substances. In intense light, plants use various mechanisms to prevent damage to their photosystems. They are able to release some light energy as heat, but
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2856-632: The excess light can also produce reactive oxygen species . While some of these can be detoxified by antioxidants , the remaining oxygen species will be detrimental to the photosystems of the plant. More specifically, the D1 subunit in the reaction center of PSII can be damaged. Studies have found that deg1 proteins are involved in the degradation of these damaged D1 subunits. New D1 subunits can then replace these damaged D1 subunits in order to allow PSII to function properly again. Wavelength In physics and mathematics , wavelength or spatial period of
2924-407: The figure at right. This change in speed upon entering a medium causes refraction , or a change in direction of waves that encounter the interface between media at an angle. For electromagnetic waves , this change in the angle of propagation is governed by Snell's law . The wave velocity in one medium not only may differ from that in another, but the velocity typically varies with wavelength. As
2992-400: The figure, wavelength is measured between consecutive corresponding points on the waveform. Localized wave packets , "bursts" of wave action where each wave packet travels as a unit, find application in many fields of physics. A wave packet has an envelope that describes the overall amplitude of the wave; within the envelope, the distance between adjacent peaks or troughs is sometimes called
3060-471: The first null of the Airy disk, to a size proportional to the wavelength of the light used, and depending on the numerical aperture : where the numerical aperture is defined as N A = n sin θ {\displaystyle \mathrm {NA} =n\sin \theta \;} for θ being the half-angle of the cone of rays accepted by the microscope objective . The angular size of
3128-417: The independent propagation of sinusoidal components. The wavelength λ of a sinusoidal waveform traveling at constant speed v {\displaystyle v} is given by where v {\displaystyle v} is called the phase speed (magnitude of the phase velocity ) of the wave and f {\displaystyle f} is the wave's frequency . In a dispersive medium ,
3196-405: The light is not altered, just where it shows up. The notion of path difference and constructive or destructive interference used above for the double-slit experiment applies as well to the display of a single slit of light intercepted on a screen. The main result of this interference is to spread out the light from the narrow slit into a broader image on the screen. This distribution of wave energy
3264-411: The local water depth. Waves that are sinusoidal in time but propagate through a medium whose properties vary with position (an inhomogeneous medium) may propagate at a velocity that varies with position, and as a result may not be sinusoidal in space. The figure at right shows an example. As the wave slows down, the wavelength gets shorter and the amplitude increases; after a place of maximum response,
3332-419: The magnesium atom. The magnesium center uniquely defines the structure as a chlorophyll molecule. The porphyrin ring of bacteriochlorophyll is saturated, and lacking alternation of double and single bonds causing variation in absorption of light. Side chains are attached to the chlorin ring of the various chlorophyll molecules. Different side chains characterize each type of chlorophyll molecule, and alters
3400-407: The medium is When wavelengths of electromagnetic radiation are quoted, the wavelength in vacuum usually is intended unless the wavelength is specifically identified as the wavelength in some other medium. In acoustics, where a medium is essential for the waves to exist, the wavelength value is given for a specified medium. The variation in speed of light with wavelength is known as dispersion , and
3468-494: The outer part of the antenna complex to the inner part. This funneling of energy is performed via resonance transfer, which occurs when energy from an excited molecule is transferred to a molecule in the ground state. This ground state molecule will be excited, and the process will continue between molecules all the way to the reaction center. At the reaction center, the electrons on the special chlorophyll molecule will be excited and ultimately transferred away by electron carriers. (If
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#17327733577453536-413: The pattern is where q is the number of slits, and g is the grating constant. The first factor, I 1 , is the single-slit result, which modulates the more rapidly varying second factor that depends upon the number of slits and their spacing. In the figure I 1 has been set to unity, a very rough approximation. The effect of interference is to redistribute the light, so the energy contained in
3604-602: The phase speed itself depends upon the frequency of the wave, making the relationship between wavelength and frequency nonlinear. In the case of electromagnetic radiation —such as light—in free space , the phase speed is the speed of light , about 3 × 10 m/s . Thus the wavelength of a 100 MHz electromagnetic (radio) wave is about: 3 × 10 m/s divided by 10 Hz = 3 m. The wavelength of visible light ranges from deep red , roughly 700 nm , to violet , roughly 400 nm (for other examples, see electromagnetic spectrum ). For sound waves in air,
3672-704: The porphyrin ring, phytol becomes the precursor of two biomarkers , pristane and phytane , which are important in the study of geochemistry and the determination of petroleum sources. The Chlorophyll a biosynthetic pathway utilizes a variety of enzymes . In most plants, chlorophyll is derived from glutamate and is synthesised along a branched pathway that is shared with heme and siroheme . The initial steps incorporate glutamic acid into 5-aminolevulinic acid (ALA); two molecules of ALA are then reduced to porphobilinogen (PBG), and four molecules of PBG are coupled, forming protoporphyrin IX. Chlorophyll synthase
3740-402: The process is termed noncyclic photophosphorylation, but if they pass through PSI and the proton pump multiple times it is called cyclic photophosphorylation. When the electron reaches photosystem I, it fills the electron deficit of light-excited reaction-center chlorophyll P700 of PSI. The electron may either continue to go through cyclic electron transport around PSI or pass, via ferredoxin, to
3808-437: The progenitors of the photosystem-containing chloroplasts of eukaryotes . Photosynthetic bacteria that cannot produce oxygen have only one photosystem, which is similar to either PSI or PSII . At the core of photosystem II is P680, a special chlorophyll to which incoming excitation energy from the antenna complex is funneled. One of the electrons of excited P680* will be transferred to a non- fluorescent molecule, which ionizes
3876-409: The range of wavelengths that can be used in photosynthesis. The addition of chlorophyll b next to chlorophyll a extends the absorption spectrum . In low light conditions, plants produce a greater ratio of chlorophyll b to chlorophyll a molecules, increasing photosynthetic yield. Absorption of light by photosynthetic pigments converts photons into chemical energy. Light energy radiating onto
3944-541: The reaction center of PSII of plants and cyanobacteria, the light energy is used to split water into oxygen, protons, and electrons. The protons will be used in proton pumping to fuel the ATP synthase at the end of an electron transport chain . A majority of the reactions occur at the D1 and D2 subunits of PSII. Both photosystem I and II are required for oxygenic photosynthesis. Oxygenic photosynthesis can be performed by plants and cyanobacteria; cyanobacteria are believed to be
4012-537: The short wavelength is associated with a high loss and the wave dies out. The analysis of differential equations of such systems is often done approximately, using the WKB method (also known as the Liouville–Green method ). The method integrates phase through space using a local wavenumber , which can be interpreted as indicating a "local wavelength" of the solution as a function of time and space. This method treats
4080-420: The special case of dispersion-free and uniform media, waves other than sinusoids propagate with unchanging shape and constant velocity. In certain circumstances, waves of unchanging shape also can occur in nonlinear media; for example, the figure shows ocean waves in shallow water that have sharper crests and flatter troughs than those of a sinusoid, typical of a cnoidal wave , a traveling wave so named because it
4148-422: The special chlorophyll a molecules in the reaction center. These special chlorophyll a molecules are located in both photosystem II and photosystem I . They are known as P680 for Photosystem II and P700 for Photosystem I. P680 and P700 are the primary electron donors to the electron transport chain. These two systems are different in their redox potentials for one-electron oxidation. The E m for P700
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#17327733577454216-424: The spectrum. Chlorophyll does not reflect light but chlorophyll-containing tissues appear green because green light is diffusively reflected by structures like cell walls. This photosynthetic pigment is essential for photosynthesis in eukaryotes , cyanobacteria and prochlorophytes because of its role as primary electron donor in the electron transport chain . Chlorophyll a also transfers resonance energy in
4284-509: The strength of the electric and the magnetic field vary. Water waves are variations in the height of a body of water. In a crystal lattice vibration , atomic positions vary. The range of wavelengths or frequencies for wave phenomena is called a spectrum . The name originated with the visible light spectrum but now can be applied to the entire electromagnetic spectrum as well as to a sound spectrum or vibration spectrum . In linear media, any wave pattern can be described in terms of
4352-529: The system locally as if it were uniform with the local properties; in particular, the local wave velocity associated with a frequency is the only thing needed to estimate the corresponding local wavenumber or wavelength. In addition, the method computes a slowly changing amplitude to satisfy other constraints of the equations or of the physical system, such as for conservation of energy in the wave. Waves in crystalline solids are not continuous, because they are composed of vibrations of discrete particles arranged in
4420-427: The wave packet, and the spread of the wavenumbers of sinusoids that make up the packet, correspond to the uncertainties in the particle's position and momentum, the product of which is bounded by Heisenberg uncertainty principle . When sinusoidal waveforms add, they may reinforce each other (constructive interference) or cancel each other (destructive interference) depending upon their relative phase. This phenomenon
4488-456: The wave vectors confined to the Brillouin zone . This indeterminacy in wavelength in solids is important in the analysis of wave phenomena such as energy bands and lattice vibrations . It is mathematically equivalent to the aliasing of a signal that is sampled at discrete intervals. The concept of wavelength is most often applied to sinusoidal, or nearly sinusoidal, waves, because in
4556-426: The wave. They are also commonly expressed in terms of wavenumber k (2π times the reciprocal of wavelength) and angular frequency ω (2π times the frequency) as: in which wavelength and wavenumber are related to velocity and frequency as: or In the second form given above, the phase ( kx − ωt ) is often generalized to ( k ⋅ r − ωt ) , by replacing the wavenumber k with a wave vector that specifies
4624-427: The wave: waves with higher frequencies have shorter wavelengths, and lower frequencies have longer wavelengths. Wavelength depends on the medium (for example, vacuum, air, or water) that a wave travels through. Examples of waves are sound waves , light , water waves and periodic electrical signals in a conductor . A sound wave is a variation in air pressure , while in light and other electromagnetic radiation
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