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Teleost

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Muscle is a soft tissue , one of the four basic types of animal tissue . Muscle tissue gives skeletal muscles the ability to contract . Muscle is formed during embryonic development , in a process known as myogenesis . Muscle tissue contains special contractile proteins called actin and myosin which interact to cause movement. Among many other muscle proteins, present are two regulatory proteins , troponin and tropomyosin .

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66-624: See text Teleostei ( / ˌ t ɛ l i ˈ ɒ s t i aɪ / ; Greek teleios "complete" + osteon "bone"), members of which are known as teleosts ( / ˈ t ɛ l i ɒ s t s , ˈ t iː l i -/ ), is, by far, the largest infraclass in the class Actinopterygii , the ray-finned fishes, and contains 96% of all extant species of fish . Teleosts are arranged into about 40 orders and 448 families . Over 26,000 species have been described. Teleosts range from giant oarfish measuring 7.6 m (25 ft) or more, and ocean sunfish weighing over 2 t (2.0 long tons; 2.2 short tons), to

132-543: A pitch accent . In Modern Greek, all vowels and consonants are short. Many vowels and diphthongs once pronounced distinctly are pronounced as /i/ ( iotacism ). Some of the stops and glides in diphthongs have become fricatives , and the pitch accent has changed to a stress accent . Many of the changes took place in the Koine Greek period. The writing system of Modern Greek, however, does not reflect all pronunciation changes. The examples below represent Attic Greek in

198-417: A batch of eggs, the male fertilises them and the larvae develop without any further parental involvement. A fair proportion of teleosts are sequential hermaphrodites , starting life as females and transitioning to males at some stage, with a few species reversing this process. A small percentage of teleosts are viviparous and some provide parental care with typically the male fish guarding a nest and fanning

264-477: A lack of contemporaneous evidence. Several theories exist about what Hellenic dialect groups may have existed between the divergence of early Greek-like speech from the common Proto-Indo-European language and the Classical period. They have the same general outline but differ in some of the detail. The only attested dialect from this period is Mycenaean Greek , but its relationship to the historical dialects and

330-419: A lesser degree. Pamphylian Greek , spoken in a small area on the southwestern coast of Anatolia and little preserved in inscriptions, may be either a fifth major dialect group, or it is Mycenaean Greek overlaid by Doric, with a non-Greek native influence. Regarding the speech of the ancient Macedonians diverse theories have been put forward, but the epigraphic activity and the archaeological discoveries in

396-797: A muscle that allows the pharyngeal jaws to have a role in grinding food in addition to transporting it. The caudal fin is homocercal , meaning the upper and lower lobes are about equal in size. The spine ends at the caudal peduncle, the base of the caudal fin, distinguishing this group from those in which the spine extends into the upper lobe of the caudal fin, such as most fish from the Paleozoic (541 to 252 million years ago). The neural arches are elongated to form uroneurals which provide support for this upper lobe. Teleosts tend to be quicker and more flexible than more basal bony fishes. Their skeletal structure has evolved towards greater lightness. While teleost bones are well calcified , they are constructed from

462-550: A prefix /e-/, called the augment . This was probably originally a separate word, meaning something like "then", added because tenses in PIE had primarily aspectual meaning. The augment is added to the indicative of the aorist, imperfect, and pluperfect, but not to any of the other forms of the aorist (no other forms of the imperfect and pluperfect exist). The two kinds of augment in Greek are syllabic and quantitative. The syllabic augment

528-399: A scaffolding of struts, rather than the dense cancellous bones of holostean fish. In addition, the lower jaw of the teleost is reduced to just three bones; the dentary , the angular bone and the articular bone . The genital and urinary tracts end behind the anus in the genital papilla ; this is observed to sex teleosts. The teleosts were first recognised as a distinct group by

594-608: A separate historical stage, though its earliest form closely resembles Attic Greek , and its latest form approaches Medieval Greek . There were several regional dialects of Ancient Greek; Attic Greek developed into Koine. Ancient Greek was a pluricentric language , divided into many dialects. The main dialect groups are Attic and Ionic , Aeolic , Arcadocypriot , and Doric , many of them with several subdivisions. Some dialects are found in standardized literary forms in literature , while others are attested only in inscriptions. There are also several historical forms. Homeric Greek

660-630: A standard subject of study in educational institutions of the Western world since the Renaissance . This article primarily contains information about the Epic and Classical periods of the language, which are the best-attested periods and considered most typical of Ancient Greek. From the Hellenistic period ( c.  300 BC ), Ancient Greek was followed by Koine Greek , which is regarded as

726-510: A vowel or /n s r/ ; final stops were lost, as in γάλα "milk", compared with γάλακτος "of milk" (genitive). Ancient Greek of the classical period also differed in both the inventory and distribution of original PIE phonemes due to numerous sound changes, notably the following: The pronunciation of Ancient Greek was very different from that of Modern Greek . Ancient Greek had long and short vowels ; many diphthongs ; double and single consonants; voiced, voiceless, and aspirated stops ; and

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792-570: Is a literary form of Archaic Greek (derived primarily from Ionic and Aeolic) used in the epic poems , the Iliad and the Odyssey , and in later poems by other authors. Homeric Greek had significant differences in grammar and pronunciation from Classical Attic and other Classical-era dialects. The origins, early form and development of the Hellenic language family are not well understood because of

858-523: Is about 1.06 kg/liter. This can be contrasted with the density of adipose tissue (fat), which is 0.9196 kg/liter. This makes muscle tissue approximately 15% denser than fat tissue. Skeletal muscle is a highly oxygen-consuming tissue, and oxidative DNA damage that is induced by reactive oxygen species tends to accumulate with age . The oxidative DNA damage 8-OHdG accumulates in heart and skeletal muscle of both mouse and rat with age. Also, DNA double-strand breaks accumulate with age in

924-418: Is added to stems beginning with consonants, and simply prefixes e (stems beginning with r , however, add er ). The quantitative augment is added to stems beginning with vowels, and involves lengthening the vowel: Some verbs augment irregularly; the most common variation is e → ei . The irregularity can be explained diachronically by the loss of s between vowels, or that of the letter w , which affected

990-666: Is called 'East Greek'. Arcadocypriot apparently descended more closely from the Mycenaean Greek of the Bronze Age. Boeotian Greek had come under a strong Northwest Greek influence, and can in some respects be considered a transitional dialect, as exemplified in the poems of the Boeotian poet Pindar who wrote in Doric with a small Aeolic admixture. Thessalian likewise had come under Northwest Greek influence, though to

1056-448: Is considered by some linguists to have been closely related to Greek . Among Indo-European branches with living descendants, Greek is often argued to have the closest genetic ties with Armenian (see also Graeco-Armenian ) and Indo-Iranian languages (see Graeco-Aryan ). Ancient Greek differs from Proto-Indo-European (PIE) and other Indo-European languages in certain ways. In phonotactics , ancient Greek words could end only in

1122-439: Is covered by a toothplate. The fourth arch is composed of pairs of ceratobranchials and epibranchials, and sometimes additionally, some pharyngobranchials and a basibranchial. The base of the lower pharyngeal jaws is formed by the fifth ceratobranchials while the second, third and fourth pharyngobranchials create the base of the upper. In the more basal teleosts the pharyngeal jaws consist of well-separated thin parts that attach to

1188-456: Is divided along the embryo 's length into somites , corresponding to the segmentation of the body (most obviously seen in the vertebral column . Each somite has three divisions, sclerotome (which forms vertebrae ), dermatome (which forms skin), and myotome (which forms muscle). The myotome is divided into two sections, the epimere and hypomere, which form epaxial and hypaxial muscles , respectively. The only epaxial muscles in humans are

1254-408: Is found within the walls of blood vessels (such smooth muscle specifically being termed vascular smooth muscle ) such as in the tunica media layer of the large ( aorta ) and small arteries , arterioles and veins . Smooth muscle is also found in lymphatic vessels, the urinary bladder , uterus (termed uterine smooth muscle ), male and female reproductive tracts , the gastrointestinal tract ,

1320-403: Is involuntary, striated muscle that is found in the walls and the histological foundation of the heart , specifically the myocardium. The cardiac muscle cells , (also called cardiomyocytes or myocardiocytes), predominantly contain only one nucleus, although populations with two to four nuclei do exist. The myocardium is the muscle tissue of the heart and forms a thick middle layer between

1386-476: Is made up of 36%. Cardiac muscle tissue is found only in the walls of the heart as myocardium , and it is an involuntary muscle controlled by the autonomic nervous system . Cardiac muscle tissue is striated like skeletal muscle, containing sarcomeres in highly regular arrangements of bundles. While skeletal muscles are arranged in regular, parallel bundles, cardiac muscle connects at branching, irregular angles known as intercalated discs . Smooth muscle tissue

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1452-454: Is non-striated and involuntary. Smooth muscle is found within the walls of organs and structures such as the esophagus , stomach , intestines , bronchi , uterus , urethra , bladder , blood vessels , and the arrector pili in the skin that control the erection of body hair. Skeletal muscle is broadly classified into two fiber types: type I (slow-twitch) and type II (fast-twitch). The density of mammalian skeletal muscle tissue

1518-399: Is non-striated. There are three types of muscle tissue in invertebrates that are based on their pattern of striation: transversely striated, obliquely striated, and smooth muscle. In arthropods there is no smooth muscle. The transversely striated type is the most similar to the skeletal muscle in vertebrates. Vertebrate skeletal muscle tissue is an elongated, striated muscle tissue, with

1584-450: Is responsible for movements of the body. Other tissues in skeletal muscle include tendons and perimysium . Smooth and cardiac muscle contract involuntarily, without conscious intervention. These muscle types may be activated both through the interaction of the central nervous system as well as by receiving innervation from peripheral plexus or endocrine (hormonal) activation. Striated or skeletal muscle only contracts voluntarily, upon

1650-409: Is voluntary muscle, anchored by tendons or sometimes by aponeuroses to bones , and is used to effect skeletal movement such as locomotion and to maintain posture . Postural control is generally maintained as an unconscious reflex, but the responsible muscles can also react to conscious control. The body mass of an average adult man is made up of 42% of skeletal muscle, and an average adult woman

1716-759: The Greek region of Macedonia during the last decades has brought to light documents, among which the first texts written in Macedonian , such as the Pella curse tablet , as Hatzopoulos and other scholars note. Based on the conclusions drawn by several studies and findings such as Pella curse tablet , Emilio Crespo and other scholars suggest that ancient Macedonian was a Northwest Doric dialect , which shares isoglosses with its neighboring Thessalian dialects spoken in northeastern Thessaly . Some have also suggested an Aeolic Greek classification. The Lesbian dialect

1782-534: The Triassic period ( Prohalecites , Pholidophorus ). However, it has been suggested that teleosts probably first evolved already during the Paleozoic era . During the Mesozoic and Cenozoic eras they diversified widely, and as a result, 96% of all living fish species are teleosts. The cladogram below shows the evolutionary relationships of the teleosts to other extant clades of bony fish, and to

1848-399: The erector spinae and small intervertebral muscles, and are innervated by the dorsal rami of the spinal nerves . All other muscles, including those of the limbs are hypaxial, and innervated by the ventral rami of the spinal nerves. During development, myoblasts (muscle progenitor cells) either remain in the somite to form muscles associated with the vertebral column or migrate out into

1914-440: The neurotransmitter acetylcholine . Smooth muscle is found in almost all organ systems such as hollow organs including the stomach , and bladder ; in tubular structures such as blood and lymph vessels , and bile ducts ; in sphincters such as in the uterus, and the eye. In addition, it plays an important role in the ducts of exocrine glands. It fulfills various tasks such as sealing orifices (e.g. pylorus, uterine os) or

1980-501: The present , future , and imperfect are imperfective in aspect; the aorist , present perfect , pluperfect and future perfect are perfective in aspect. Most tenses display all four moods and three voices, although there is no future subjunctive or imperative. Also, there is no imperfect subjunctive, optative or imperative. The infinitives and participles correspond to the finite combinations of tense, aspect, and voice. The indicative of past tenses adds (conceptually, at least)

2046-434: The respiratory tract , the arrector pili of skin , the ciliary muscle , and the iris of the eye . The structure and function is basically the same in smooth muscle cells in different organs, but the inducing stimuli differ substantially, in order to perform individual actions in the body at individual times. In addition, the glomeruli of the kidneys contain smooth muscle-like cells called mesangial cells . Cardiac muscle

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2112-1031: The 5th century BC. Ancient pronunciation cannot be reconstructed with certainty, but Greek from the period is well documented, and there is little disagreement among linguists as to the general nature of the sounds that the letters represent. /oː/ raised to [uː] , probably by the 4th century BC. Greek, like all of the older Indo-European languages , is highly inflected. It is highly archaic in its preservation of Proto-Indo-European forms. In ancient Greek, nouns (including proper nouns) have five cases ( nominative , genitive , dative , accusative , and vocative ), three genders ( masculine , feminine , and neuter ), and three numbers (singular, dual , and plural ). Verbs have four moods ( indicative , imperative , subjunctive , and optative ) and three voices (active, middle, and passive ), as well as three persons (first, second, and third) and various other forms. Verbs are conjugated through seven combinations of tenses and aspect (generally simply called "tenses"):

2178-495: The Archaic period of ancient Greek (see Homeric Greek for more details): Μῆνιν ἄειδε, θεά, Πηληϊάδεω Ἀχιλῆος οὐλομένην, ἣ μυρί' Ἀχαιοῖς ἄλγε' ἔθηκε, πολλὰς δ' ἰφθίμους ψυχὰς Ἄϊδι προΐαψεν ἡρώων, αὐτοὺς δὲ ἑλώρια τεῦχε κύνεσσιν οἰωνοῖσί τε πᾶσι· Διὸς δ' ἐτελείετο βουλή· ἐξ οὗ δὴ τὰ πρῶτα διαστήτην ἐρίσαντε Ἀτρεΐδης τε ἄναξ ἀνδρῶν καὶ δῖος Ἀχιλλεύς. The beginning of Apology by Plato exemplifies Attic Greek from

2244-642: The Classical period of ancient Greek. (The second line is the IPA , the third is transliterated into the Latin alphabet using a modern version of the Erasmian scheme .) Ὅτι [hóti Hóti μὲν men mèn ὑμεῖς, hyːmêːs hūmeîs,   Muscle Muscle tissue varies with function and location in the body. In vertebrates , the three types are: Skeletal muscle tissue consists of elongated, multinucleate muscle cells called muscle fibers , and

2310-537: The German ichthyologist Johannes Peter Müller in 1845. The name is from Greek teleios , "complete" + osteon , "bone". Müller based this classification on certain soft tissue characteristics, which would prove to be problematic, as it did not take into account the distinguishing features of fossil teleosts. In 1966, Greenwood et al. provided a more solid classification. The oldest fossils of teleosteomorphs (the stem group from which teleosts later evolved) date back to

2376-550: The aorist. Following Homer 's practice, the augment is sometimes not made in poetry , especially epic poetry. The augment sometimes substitutes for reduplication; see below. Almost all forms of the perfect, pluperfect, and future perfect reduplicate the initial syllable of the verb stem. (A few irregular forms of perfect do not reduplicate, whereas a handful of irregular aorists reduplicate.) The three types of reduplication are: Irregular duplication can be understood diachronically. For example, lambanō (root lab ) has

2442-419: The augment when it was word-initial. In verbs with a preposition as a prefix, the augment is placed not at the start of the word, but between the preposition and the original verb. For example, προσ(-)βάλλω (I attack) goes to προσ έ βαλoν in the aorist. However compound verbs consisting of a prefix that is not a preposition retain the augment at the start of the word: αὐτο(-)μολῶ goes to ηὐ τομόλησα in

2508-469: The body to form all other muscles. Myoblast migration is preceded by the formation of connective tissue frameworks, usually formed from the somatic lateral plate mesoderm . Myoblasts follow chemical signals to the appropriate locations, where they fuse into elongate skeletal muscle cells. The primary function of muscle tissue is contraction . The three types of muscle tissue (skeletal, cardiac and smooth) have significant differences. However, all three use

2574-438: The center of Greek scholarship, this division of people and language is quite similar to the results of modern archaeological-linguistic investigation. One standard formulation for the dialects is: West vs. non-West Greek is the strongest-marked and earliest division, with non-West in subsets of Ionic-Attic (or Attic-Ionic) and Aeolic vs. Arcadocypriot, or Aeolic and Arcado-Cypriot vs. Ionic-Attic. Often non-West

2640-615: The dialect of Sparta ), and Northern Peloponnesus Doric (including Corinthian ). All the groups were represented by colonies beyond Greece proper as well, and these colonies generally developed local characteristics, often under the influence of settlers or neighbors speaking different Greek dialects. After the conquests of Alexander the Great in the late 4th century BC, a new international dialect known as Koine or Common Greek developed, largely based on Attic Greek , but with influence from other dialects. This dialect slowly replaced most of

2706-435: The eggs to keep them well-oxygenated. Teleosts are economically important to humans, as is shown by their depiction in art over the centuries. The fishing industry harvests them for food, and anglers attempt to capture them for sport . Some species are farmed commercially, and this method of production is likely to be increasingly important in the future. Others are kept in aquariums or used in research, especially in

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2772-680: The exact location of the muscle. Sub-categorization of muscle tissue is also possible, depending on among other things the content of myoglobin , mitochondria , and myosin ATPase etc. The word muscle comes from Latin musculus , diminutive of mus meaning mouse , because the appearance of the flexed biceps resembles the back of a mouse. The same phenomenon occurred in Greek , in which μῦς, mȳs , means both "mouse" and "muscle". There are three types of muscle tissue in vertebrates: skeletal , cardiac , and smooth . Skeletal and cardiac muscle are types of striated muscle tissue . Smooth muscle

2838-405: The fibres ranging from 3-8 micrometers in width and from 18 to 200 micrometers in breadth. In the uterine wall, during pregnancy, they enlarge in length from 70 to 500 micrometers. Skeletal striated muscle tissue is arranged in regular, parallel bundles of myofibrils , which contain many contractile units known as sarcomeres , which give the tissue its striated (striped) appearance. Skeletal muscle

2904-409: The fields of genetics and developmental biology . Distinguishing features of the teleosts are mobile premaxilla , elongated neural arches at the end of the caudal fin and unpaired basibranchial toothplates. The premaxilla is unattached to the neurocranium (braincase); it plays a role in protruding the mouth and creating a circular opening. This lowers the pressure inside the mouth, sucking

2970-673: The forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek ( c.  1400–1200 BC ), Dark Ages ( c.  1200–800 BC ), the Archaic or Epic period ( c.  800–500 BC ), and the Classical period ( c.  500–300 BC ). Ancient Greek was the language of Homer and of fifth-century Athenian historians, playwrights, and philosophers . It has contributed many words to English vocabulary and has been

3036-718: The four-limbed vertebrates ( tetrapods ) that evolved from a related group of bony fish during the Devonian period . Approximate divergence dates (in millions of years, mya ) are from Near et al., 2012. Coelacanths [REDACTED] Lungfish [REDACTED] Lissamphibia [REDACTED] Mammals [REDACTED] Sauropsida ( reptiles , birds ) [REDACTED] Polypteriformes ( bichirs , reedfishes ) [REDACTED] Acipenseriformes ( sturgeons , paddlefishes ) [REDACTED] Lepisosteiformes ( gars ) [REDACTED] Amiiformes ( bowfin ) [REDACTED] Teleostei [REDACTED] The phylogeny of

3102-561: The historical Dorians . The invasion is known to have displaced population to the later Attic-Ionic regions, who regarded themselves as descendants of the population displaced by or contending with the Dorians. The Greeks of this period believed there were three major divisions of all Greek people – Dorians, Aeolians, and Ionians (including Athenians), each with their own defining and distinctive dialects. Allowing for their oversight of Arcadian, an obscure mountain dialect, and Cypriot, far from

3168-476: The historical circumstances of the times imply that the overall groups already existed in some form. Scholars assume that major Ancient Greek period dialect groups developed not later than 1120 BC, at the time of the Dorian invasions —and that their first appearances as precise alphabetic writing began in the 8th century BC. The invasion would not be "Dorian" unless the invaders had some cultural relationship to

3234-466: The influence of the central nervous system. Reflexes are a form of non-conscious activation of skeletal muscles, but nonetheless arise through activation of the central nervous system, albeit not engaging cortical structures until after the contraction has occurred. The different muscle types vary in their response to neurotransmitters and hormones such as acetylcholine , noradrenaline , adrenaline , and nitric oxide depending on muscle type and

3300-440: The jaw musculature which make it possible for them to protrude their jaws outwards from the mouth . This is of great advantage, enabling them to grab prey and draw it into the mouth . In more derived teleosts, the enlarged premaxilla is the main tooth-bearing bone, and the maxilla, which is attached to the lower jaw, acts as a lever, pushing and pulling the premaxilla as the mouth is opened and closed. Other bones further back in

3366-446: The minute male anglerfish Photocorynus spiniceps , just 6.2 mm (0.24 in) long. Including not only torpedo-shaped fish built for speed, teleosts can be flattened vertically or horizontally, be elongated cylinders or take specialised shapes as in anglerfish and seahorses . The difference between teleosts and other bony fish lies mainly in their jaw bones; teleosts have a movable premaxilla and corresponding modifications in

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3432-407: The mouth serve to grind and swallow food. Another difference is that the upper and lower lobes of the tail (caudal) fin are about equal in size. The spine ends at the caudal peduncle , distinguishing this group from other fish in which the spine extends into the upper lobe of the tail fin. Teleosts have adopted a range of reproductive strategies . Most use external fertilisation: the female lays

3498-430: The movement of actin against myosin to create contraction. In skeletal muscle, contraction is stimulated by electrical impulses transmitted by the motor nerves . Cardiac and smooth muscle contractions are stimulated by internal pacemaker cells which regularly contract, and propagate contractions to other muscle cells they are in contact with. All skeletal muscle and many smooth muscle contractions are facilitated by

3564-406: The neurocranium, pectoral girdle , and hyoid bar . Their function is limited to merely transporting food, and they rely mostly on lower pharyngeal jaw activity. In more derived teleosts the jaws are more powerful, with left and right ceratobranchials fusing to become one lower jaw; the pharyngobranchials fuse to create a large upper jaw that articulates with the neurocranium. They have also developed

3630-468: The nodes (so, the pattern of branching shown is likely to be correct). They calibrated (set actual values for) branching times in this tree from 36 reliable measurements of absolute time from the fossil record. The teleosts are divided into the major clades shown on the cladogram, with dates, following Near et al. More recent research divide the teleosts into two major groups: Eloposteoglossocephala (Elopomorpha + Osteoglossomorpha) and Clupeocephala (the rest of

3696-508: The older dialects, although the Doric dialect has survived in the Tsakonian language , which is spoken in the region of modern Sparta. Doric has also passed down its aorist terminations into most verbs of Demotic Greek . By about the 6th century AD, the Koine had slowly metamorphosed into Medieval Greek . Phrygian is an extinct Indo-European language of West and Central Anatolia , which

3762-660: The outer epicardium layer and the inner endocardium layer. Coordinated contractions of cardiac muscle cells in the heart propel blood out of the atria and ventricles to the blood vessels of the left/body/systemic and right/lungs/pulmonary circulatory systems . This complex mechanism illustrates systole of the heart. Cardiac muscle cells, unlike most other tissues in the body, rely on an available blood and electrical supply to deliver oxygen and nutrients and to remove waste products such as carbon dioxide . The coronary arteries help fulfill this function. All muscles are derived from paraxial mesoderm . The paraxial mesoderm

3828-487: The perfect stem eilēpha (not * lelēpha ) because it was originally slambanō , with perfect seslēpha , becoming eilēpha through compensatory lengthening. Reduplication is also visible in the present tense stems of certain verbs. These stems add a syllable consisting of the root's initial consonant followed by i . A nasal stop appears after the reduplication in some verbs. The earliest extant examples of ancient Greek writing ( c.  1450 BC ) are in

3894-436: The prey inside. The lower jaw and maxilla are then pulled back to close the mouth, and the fish is able to grasp the prey . By contrast, mere closure of the jaws would risk pushing food out of the mouth. In more advanced teleosts, the premaxilla is enlarged and has teeth, while the maxilla is toothless. The maxilla functions to push both the premaxilla and the lower jaw forward. To open the mouth, an adductor muscle pulls back

3960-524: The skeletal muscle of mice. Smooth muscle is involuntary and non-striated. It is divided into two subgroups: the single-unit (unitary) and multiunit smooth muscle . Within single-unit cells, the whole bundle or sheet contracts as a syncytium (i.e. a multinucleate mass of cytoplasm that is not separated into cells). Multiunit smooth muscle tissues innervate individual cells; as such, they allow for fine control and gradual responses, much like motor unit recruitment in skeletal muscle. Smooth muscle

4026-517: The syllabic script Linear B . Beginning in the 8th century BC, however, the Greek alphabet became standard, albeit with some variation among dialects. Early texts are written in boustrophedon style, but left-to-right became standard during the classic period. Modern editions of ancient Greek texts are usually written with accents and breathing marks , interword spacing , modern punctuation , and sometimes mixed case , but these were all introduced later. The beginning of Homer 's Iliad exemplifies

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4092-471: The teleosts has been subject to long debate, without consensus on either their phylogeny or the timing of the emergence of the major groups before the application of modern DNA -based cladistic analysis. Near et al. (2012) explored the phylogeny and divergence times of every major lineage, analysing the DNA sequences of 9 unlinked genes in 232 species. They obtained well-resolved phylogenies with strong support for

4158-1574: The teleosts). Hiodontiformes ( mooneyes ) [REDACTED] Osteoglossiformes ( bonytongues , elephantfishes ) [REDACTED] Elopiformes ( tenpounders , tarpons ) [REDACTED] Albuliformes ( Japanese gissus and bonefishes ) [REDACTED] Notacanthiformes (deep sea spiny eels) [REDACTED] Anguilliformes (true eels ) [REDACTED] Clupeiformes ( herrings ) [REDACTED] Alepocephaliformes ( slickheads ) [REDACTED] Gonorynchiformes ( milkfish ) [REDACTED] Cypriniformes ( minnows , carps , loaches ) [REDACTED] Characiformes ( tetras , piranhas ) [REDACTED] Gymnotiformes (knifefish and electric eels ) [REDACTED] Siluriformes (catfish) [REDACTED] Lepidogalaxiiformes (salamanderfish) Argentiniformes (marine smelts) [REDACTED] Galaxiiformes ( whitebait , mudfishes) [REDACTED] Esociformes ( pike ) [REDACTED] Salmoniformes ( salmon , trout ) [REDACTED] Stomiiformes (dragonfish) [REDACTED] Osmeriformes ( smelt ) [REDACTED] Ateleopodiformes (jellynoses) [REDACTED] Aulopiformes (lizardfish) [REDACTED] Myctophiformes ( lanternfish ) [REDACTED] Lampriformes ( oarfish , opah , ribbonfish ) [REDACTED] Percopsiformes (troutperches) [REDACTED] Zeiformes (dories) [REDACTED] Stylephoriformes (tube-eyes/thread-fins) Ancient Greek Ancient Greek ( Ἑλληνῐκή , Hellēnikḗ ; [hellɛːnikɛ́ː] ) includes

4224-508: The top of the maxilla, pushing the lower jaw forward. In addition, the maxilla rotates slightly, which pushes forward a bony process that interlocks with the premaxilla. The pharyngeal jaws of teleosts, a second set of jaws contained within the throat, are composed of five branchial arches , loops of bone which support the gills . The first three arches include a single basibranchial surrounded by two hypobranchials, ceratobranchials, epibranchials and pharyngobranchials. The median basibranchial

4290-426: The transport of the chyme through wavelike contractions of the intestinal tube. Smooth muscle cells contract more slowly than skeletal muscle cells, but they are stronger, more sustained and require less energy. Smooth muscle is also involuntary, unlike skeletal muscle, which requires a stimulus. Cardiac muscle is the muscle of the heart. It is self-contracting, autonomically regulated and must continue to contract in

4356-480: Was Aeolic. For example, fragments of the works of the poet Sappho from the island of Lesbos are in Aeolian. Most of the dialect sub-groups listed above had further subdivisions, generally equivalent to a city-state and its surrounding territory, or to an island. Doric notably had several intermediate divisions as well, into Island Doric (including Cretan Doric ), Southern Peloponnesus Doric (including Laconian ,

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