Sclerocytes secrete the mineral elements of the skeleton - spicules that perform a supporting function and maintain the shape of the body of the sponge. In silicon sponges, sclerocytes carry out intracellular synthesis of spicules. Of all Metazoa, only ordinary and glass sponges use silicon , not calcium , to build the skeleton. By the size of the formed spicules, megasclerocyte and microsclerocytes are secreted. A spicule is synthesized inside a vacuole formed by a special membrane - silicallemma, which does not contact any cell membranes [59] . Sclerocytes of calcareous sponges originate from pinacocytes and, possibly, choanocytes [60] .
In addition to the internal main skeleton, many sponges also have a skin (ectosomal) skeleton that protects the sponge from predators and other sponges. Its structure is diverse and can include both horizontally oriented spicules collected in piles and vertically oriented spicules forming a palisade [61] .
Spongocytes form a periospicular spongin or “horn” skeleton in ordinary sponges lacking a mineral skeleton. These cells migrate to sclerocytes synthesizing spicules or ready-made spicules and attach sponging molecules to them - a special form of collagen containing a high concentration of iodine . Spongin is found only in sponges [62] and has the appearance of thick skeletal fibers [63] . It is believed that spongocytes are characterized by positive chemotaxis to silicon. With the formation of the organic skeleton, spongocytes are grouped by connecting finger-shaped contacts. In ordinary sponges, spongocytes, like sclerocytes, are believed to be derivatives of the archaeocytic line [64] .
Secretory Protective Cells
All cells of this group can be divided into amoeboid cells and cells with inclusions. The main functions of this group of cells are protective (phagocytic, bactericidal ), storage, secretion of the main substance of mesochil, as well as the transfer of particles of food and oxygen [64] .
- Amoeboid cells
Ordinary sponges have typical amoeboid cells - archeocytes, which are distinguished by a large nucleus and nucleolus, as well as numerous large lysosomes [57] capable of phagocytosis . Homoscleromorpha does not have such cells in all species. Currently, archaeocytes are understood as totipotent ( stem ) cells, like any cells that have a nucleus with a nucleolus and numerous phagosomes . Archaeocytes are the most actively dividing fraction of free cells. They are actively involved in asexual and sexual reproduction, as well as regenerative morphogenesis [65] .
Bacteriocytes are motile amoeboid cells with vacuoles in which prokaryotic endosymbionts live. So far, they have been identified only in some types of ordinary sponges. Some bacteriocytes contain inside one large vacuole with bacteria, while others have numerous small vacuoles in which one or more bacteria are enclosed. Bacteriocytes can be quite large (up to 20 microns). In some species, bacteriocytes carry out a vertical transfer of prokaryotic symbionts from the mother to the daughter, penetrating the embryo. Bacteriocytes are believed to come from the archaeocytic line [66] .
- Cells with inclusions
This is a heterogeneous group of cells containing special inclusions in the cytoplasm. Unlike archeocytes, their rough endoplasmic reticulum is reduced, there are no phagosomes, acid phosphatase activity is not detected, and there is no nucleolus in the nuclei. According to the size of inclusions, cells containing large inclusions and cells containing small inclusions can be distinguished. The first group includes:
- vacuolar cells containing one or more electron-transparent or light vacuoles;
- globular cells containing a little free cytoplasm, one or two large globules of unknown composition and a small deformed nucleus;
- spherical cells are filled with large rounded inclusions of various nature and chemical composition. Their entire cytoplasm is filled with large inclusions limited by membranes (spherules). Usually spherical cells are localized in the ectosome along the channels of the aquifer system. Their functions are diverse: they contain toxic metabolites , participate in excretion, accumulate lectins , participate in the maintenance of the extracellular matrix;
- granular cells differ from spherical cells only in a smaller size of inclusions [67] .
The second group includes:
- gray cells, or glycocytes. They were found in many ordinary sponges and contain numerous ovoid inclusions limited by membranes, as well as glycogen rosettes. Their main role is in glycogen metabolism and collagen secretion , together with archaeocytes they are involved in various morphogenesis;
- microgranular cells have a cytoplasm filled with small electron-dense cells. Their functions have been little studied, but it is known for certain that they do not participate in collagen synthesis [68] .
Move cells
The ability of most cells composing the body to move freely is a hallmark of sponges. Mesochilus cells move almost constantly. During the restructuring of the aquifer, endopinacocytes and choanocytes can change their localization, as a result of which new flagellar chambers form, new canal branches merge and form. This ensures the optimization of the flow of water in the body of the sponge as it grows or in connection with changes in the environment. Independent and frequent movements of cells in the bodies of the sponges are possible due to the absence of real intercellular contacts and the basal lamina in most of them [69] .
Sintzitiy glass sponges
water flow spicules primary syncytium choanocytic cells
Glass sponges do not have typical choanoderma or pinacoderm. Their living tissue is a three-dimensional network, similar to a web, which is formed by numerous strands. There are no membranes separating the cells, and the cytoplasm is continuous within the tissue; therefore, the tissue of glass sponges is called trabecular syncytium or the trabecular network. Instead of the choanoderma, there is another syncytium - choanosyntsium, and instead of the typical choanocytes - collar bodies, each of which has a collar and flagellum, but no core. Collar bodies are collected in groups in separate pockets surrounded by a trabecular meshwork. In fact, the collar bodies are the outgrowths of a single stem cell (choanoblast) containing the nucleus. Mesochilus extends inside each strand of the trabecular network. It contains spicules, collagen fibers, and individual cells: sclerocytes, archeocytes, and, most likely, germ cells [70] .
Physiology
Locomotion and body shape change
The vast majority of sponges are sedentary attached organisms, however, some species are capable of limited locomotion . For example, the freshwater sponge Ephydatia , as well as some marine species, can move along the substrate at a speed of 1 to 4 mm per day. Such a movement is achieved as a result of the amoeboid movement of pinacocytes and other types of cells, which occurs to some extent in a coordinated manner. Some species, such as Clathrina coriceae , can contract and change the diameter of the oskulums with the help of myocytes. Thanks to these mechanisms, sponges can temporarily stop the flow of water through their aquifer system, for example, with an increase in the amount of suspension in water [71] . Adult sponges of all four classes can expel particles of substrate from themselves by contraction [72] .
It is shown that changes in body volume and shape can also be endogenous and have a daily rhythm. It is not known exactly what is responsible for signal transmission between cells in the absence of a nervous system : cell conductivity, chemical diffusion, or electrical conductivity. Apparently, any of the options can take place, since bioelectric cell potentials and neurotransmitters are revealed in the sponges. Moreover, in their genomes there are genes orthologous to the genes involved in the functioning of the neurosensory system of eumetasoids [73] . However, due to the lack of specialized intercellular contacts, the signal in the body of the sponge propagates very slowly. An exception is glass sponges with a syncytial structure. For example, in Rhabdocalyptus dawsoni action potentials quickly spread along syncytial cords, as a result of which the beating of flagella and the flow of water through the body stop [74] .
Nutrition
Most sponges filter out small food particles from the water that flows through their body. Particle sizes typically range from 50 to 1 μm, but may be even smaller. Unicellular planktonic organisms, for example, dinoflagellates and bacteria, viruses , small organic detritus, and possibly even organic compounds dissolved in water, fall into these limits [75] .
Sponges do not have a population of cells specialized in digestive function. Almost all integumentary cells and many cells of the inner space of the sponge are involved in the capture of food particles. It has been shown that choanocytes, archeocytes, exopinacocytes, endopinacocytes and basopinacocytes have phagocytic and digestive activity. Food grabbing can also occur in prosopiles and ducts [36] . Phagocytic activity was detected in myocytes, secretory globular cells, sclerocytes, lofocytes. Small particles can penetrate the mesochil between exopinacocytes, however, the mechanism of this movement is unclear. Some mesochil cells can amoeboid extend their processes into the cavity of the adduction canals between pinacocytes and carry out phagocytosis. However, in most cases, bacteria and other picoplankton are captured by pseudopodial outgrowths of choanocytes [36] .
Hoanocytes can capture food particles and the basal part, and apical (in the collar). The choanocytes and pinacocytes that phagocytosed food particles either transmit them to archaeocytes or excrete them into the mesochil by exocytosis . Archaeocytes and choanocytes digest food in the , and archaeocytes are also likely to store nutrients (glycogen and lipids ). Particularly large particles settling in the channels are absorbed by endopinacocytes and archaeocytes, which penetrate the channels through the pinakocyte layer or through porocytes. Enzymes of amylase , lipase , protease , esterase [76] [77] take part in the breakdown of food in sponges.
Among the sponges, carnivorous species are also known - ordinary sponges of the families , as well as some species of the families Guitarridae and Esperiopsidae. Only 137 species of predatory sponges are known [78] . Representatives of Cladorhizidae feed on crustaceans and other small animals that adhere to long sticky cell-like threads spread out by a sponge. When the prey sticks, the thread is shortened, pulling the victim to the body of the sponge. Next, the sponge slowly envelops the victim and digests it, most likely with the help of archaeocytes. There is no evidence that the sponge uses poison or toxins . Some predatory sponges use hook-shaped spicules to catch prey [79] . Predatory sponges are devoid of choanocytes and the aquifer system [80] [77] , although sponges of the genus use their modified aquifer system to suck prey into the body. Most predatory sponges are inhabitants of great depths (up to 8840 m) [79] [81] .
Undigested food residues, as well as inorganic particles that have fallen into the sponge with a stream of water, are subject to removal from the body of the sponge. The latter must be removed: they can clog the drive channels and interfere with their operation. The particle stuck in the lead canal is phagocytosed by the archeocyte and then carries it closer to the exit from the aquifer system, releasing exocytosis into the discharge canal. In sponges using foreign particles to construct the skeleton, the captured particle can be transferred to the place of synthesis of skeleton elements [82] .
Inland transport, gas exchange and emissions
The entry of gases and metabolic products (mainly ammonia ) from the body into the water flowing through the aquifer system provides simple diffusion . It may play a role in the distribution of nutrients throughout the body of the sponge, although archaeocytes are mainly involved. Species of the genus have special internal fibers along which, as on rails, archaeocytes filled with nutrients move [83] .
In glass sponges, transport is carried out within syncytium. After the food particles have been captured by the collar bodies and loaded into vacuoles, vacuoles are moved throughout the syncytium along the microtubules using the dynein motor protein . In the same way, vesicles move in the pseudopodia of foraminifera and in the axons of neurons of some higher animals [83] .
Since there are practically no intercellular contacts in the pinacoderm and choanoderm, these cell layers do not form a reliable barrier between the mesochil and water; therefore, water constantly enters the body of the sponges. The cells of freshwater sponges usually have contractile vacuoles that carry out osmoregulation at the level of individual cells, but not the entire body of the animal [84] .
Regulation of water flow
Sponges have various devices for regulating the flow of water through the aquifer system. In glass sponges, flagella cease to break if particles of bottom sediment fall into the sponge or if its tissue is damaged. In this case, the action potential spreads along the syncytial tissue, and the flagella resume beating when their membrane potential reaches a normal value. It is possible that the action potential causes calcium to enter the collar bodies, and the beating of the flagellum is restored when calcium is pumped out of the cell [36] .
In sponges with the cellular structure of tissues, the action potential, as far as is known, cannot spread throughout the body. However, in response to poor water quality, physical or chemical irritation, or even in a spontaneous rhythm, such sponges can slowly reduce the channels of their aquifer system, close the ostia and oskulum, as a result of which the water flow slows down or stops altogether. At the same time, the flagella of choanocytes do not stop beating and work continuously, although in some choanocytes the flagella beat slower than in others. The beating of flagella occurs along a sinusoid and in one plane, and within the same choanocyte chamber the flagella of each choanocyte beat in their own plane; however, in response to the damaging effects (rupture of the sponge, lack of oxygen), some flagella slow their beating and may even stop for several hours. Currently, it is believed that only glass sponges are capable of instantly stopping the beating of flagella due to their syncytial structure. A glass sponge Oopsacas minuta can contract its body very slowly, with contractions caused by calcium waves traveling through syncytium. With the contraction of the body of the sponge, the choanocyte chambers and individual choanocytes also contract [36] .
Immune System
Sponges have an effective system of innate immunity , which allows sponges to distinguish between prokaryotic pathogens , prokaryotic endosymbionts and prokaryotes serving as food. It has been shown that when spongy bacteria enter the sponge, they produce special substances, for example, in the case of the sponge , which suppress the immune response and apoptosis . In response to these molecules, the sponge begins to synthesize endocytosis proteins abundantly, probably to control the bacterial population. Some membrane proteins have eukaryotic domains responsible for protein-protein interactions with the eukaryotic extracellular matrix and intercellular adhesion , which probably plays a role in recognition. Pathogenic gram-positive and gram-negative bacteria carry PAMP], which are recognized by sponge cells. For example, lipopolysaccharides (LPS), which are PAMP gram-negative bacteria, are recognized in sponges, as in other Metazoa , by Toll-like receptors that trigger an immune response [85] .
Sponges reject grafts from other species, but accept grafts from individuals of their species. In some sea sponges, gray cells play a key role in the rejection of foreign material. When a foreign fragment is introduced, gray cells release chemicals that inhibit the movement of cells in the affected area so that cells of the foreign fragment cannot use the transport system of the sponge. After that, the gray cells are grouped and release toxins that kill all cells in the affected area. Such an activated state of the immune system can last up to three weeks [86] .
Reproduction
Sponges have both sexual and asexual reproduction. Sponges can be oviparous and viviparous . Ovarian sponges are usually dioecious, and in the second case they are often hermaphrodites [87] . However, there is a known case of a sponge changing its sex during the year [88] .
Sexual reproduction
There are no gonads in the sponges, and gamete formation is not confined to a specific site of the mesochil, so gametes are evenly distributed over it, but they are always located at a distance covered by diffusion from the channel or chamber [89] . Sex cells are formed by direct transformation from somatic cells : nucleolar amoebocytes and choanocytes, but the main stages of gamete development in sponges are similar to those in other animals [87] .
With the formation of male gametes ( spermatogenesis ) in glass and ordinary sponges, as well as Homoscleromorpha , temporary spherical formations are formed, surrounded by flattened somatic cells - spermatocysts, in which spermatogenesis proceeds. In calcareous sponges, spermatocysts are not formed. Within the spermatocyst, male germ cells develop synchronously, but in Homoscleromorpha, within the spermatocyst, a sperm maturation gradient is observed, which also brings this group of sponges closer to the eumetases. Most sponges have primitive sperm and have a large cytoplasm with several mitochondria ; acrosomes are usually not. However, it was found by electron microscopy that the spermatozoa of all Homoscleromorpha and some common sponges have acrosomes. In calcareous sponges, sperm are atypical; they have a spherical shape and lack a flagellum [90] .
Developing female gametes are located diffusely or in small groups in the choanosome. In most sponges, the typical formation of yolk does not occur in oogenesis; instead, various forms of oocyte transmission from auxiliary cells of the nutrient material from which the secondary yolk is formed predominate. In this case, phagocytosis of whole cells or their fragments entering the phagolysosomes can occur. The egg and auxiliary cells are sometimes enclosed in a special follicle that has cell walls [89] . In Homoscleromorpha and egg-laying ordinary sponges, the yolk is formed due to endogenous synthesis. In oocytes, ooplasm is not segregated. Oviparous sponges do not have egg membranes, but in oviparous ordinary sponges a primitive vitelline membrane is formed during oogenesis [91] .
During the breeding season, the wall of the spermatocyst is destroyed, and the sperm enter the outlet channels (less often, the atrium) and are thrown out through the oskulums. Some tropical sponges suddenly eject milky clouds of sperm from the oskulums. When the ejected spermatozoa approach another sponge, then, together with the currents of water, they are transferred to the aquifer system. Inside the sponge, they approach the surface of the choanoderm or in the flagellum chambers, where they are phagocytized, but not digested, choanocytes. A choanocyte that has absorbed a sperm cell loses its flagellum and collar and turns into an amoeboid cell (a carrier cell), which transfers the sperm nucleus to the egg cell and either transfers the nucleus to it or is phagocytized by the egg [89] . Some sponges release their eggs, along with auxiliary cells, into the water. Often sponges throw eggs and sperm into the water in synchronization; this is called "smoking" or "boiling" of the sponges. In the tropical reef sponge synchronization of sperm and egg release is very clear, so that once a year and three days of smoking can be predicted in advance [92] .
Asexual Reproduction
Asexual reproduction is described in all classes of sponges and can be carried out by fragmentation, hemmulogenesis, and budding [93] .
Fragmentation is considered the most primitive way of asexual reproduction. It can occur during severe storms, due to predators (fish, turtles), infectious diseases, rupture or cracking of the substrate, as a result of which the body of the sponge breaks up into fragments. Separated pieces survive due to the ability of the sponges to regenerate (see below for more details). They attach to the substrate, rearrange, and turn into a new sponge [94] . Since fragmentation is not a regular stage in the development of a sponge, but an event caused by random external factors, no cytological changes occur during fragmentation. Fragmentation is most characteristic of ordinary sponges [95] .
Budding is a rather rare way of reproducing sponges: as an obligate one, it appears in the life cycle only in representatives of two families ( and ), although it is presented on many occasions in many families [96] . Moreover, in most sponges (with the exception of Homoscleromorpha and Radiospongilla cerebellata ), a dense accumulation of cells, mainly archeocytes, forms on the body of the mother sponge. The formation of such a kidney is based on the migration of pluripotent cells with their subsequent differentiation. This kidney is deprived of choanocytic chambers, channels, oskulum and after separation from the maternal sponge is attached to the substrate; there it begins to grow and form an aquifer system. When settling, the kidneys can merge with each other, as a result of which larger sponges develop. In Clathrina, the kidneys form as the swelling of the ends of the tubes that represent the body of the sponge. In some Tethya species , the buds form on special stems, while in Oscarella and Aplysilla they are papillas that separate from the parent organism and give rise to new sponges [94] . In Homoscleromorpha, the mechanism of formation of a new sponge is completely different: in these sponges, the kidneys are the outgrowths of the body of the parent sponge, and proliferation does not occur during their formation, and the cellular composition of the kidney is identical to the cellular composition of the parent sponge [97] .
Hemmulogenesis is the most complex type of asexual reproduction of sponges. Its result is the formation of gemmules - resting stages, designed to survive adverse conditions, as well as the spread of sponges. Gemmules form freshwater and some marine sponges in the amount of hundreds and thousands [94] . Outside, the gemmula is covered with a non-cellular protective membrane, and inside it is a mass of pluripotent cells and tezocytes (inactive archaeocytes) filled with a reserve substance [98] . Periodically, the body of the sponge dies and completely disintegrates, so that only gemmules remain [99] .
Gemmules are extremely resistant to many adverse factors and can germinate even after two months of maintenance at −80 ° C and even −100 ° C. A quarter of gemmules germinate after four months of dehydration at 5 ° C [100] . Sponge specialists often store moistened gemmules in the refrigerator and germinate them for experiments or observations [101] .
For many species whose life cycle includes asexual reproduction, the alternation of sexual and asexual reproduction is characteristic. In particular, within one sponge, the formation of gemmules does not begin earlier than the formation of yolk in oocytes due to competition for cellular material. This is also the reason for the decrease in the activity of sexual reproduction during asexual reproduction. In many freshwater sponges, asexual reproduction intensifies after sexual reproduction and completes the active phase of the life cycle in sponge populations living in unstable conditions [102] .
The ability to regenerate sponges is superior to all other Metazoa. In case of damage to the body of the sponge, most of the archeocytes turn into pinacocytes and choanocytes, restoring the pinacoderm and choanoderm; The whole process takes several days. The ability to regenerate is actively used in aquaculture of a toilet sponge, when the sponge is cut into pieces and after a few months full sponges develop from the pieces. In addition, a high ability to regenerate sponges is used by researchers studying their physiology. A piece of sponge is placed on a coverslip, to which the sponge grows, and then part of the sponge is covered with a second coverslip. As a result, a very thin layer of living matter is formed, which is convenient to study under a microscope. In the study of individual cells of the sponge, it is divided into individual cells by physical or chemical means. If one of these cells is put into a vessel, then one young sponge will develop at the bottom of the vessel [103] . Interestingly, sponge cells are able to re-react to form a normal sponge after dividing a sponge fragment into individual cells. An intermediate stage of this process is the formation of primmorphs - clusters of undifferentiated cells coated with exopinacoderm. Further, the cells undergo transdifferentiation with the formation of a normal sponge, and the fate of the cell during transdifferentiation is determined by its position in the multicellular aggregate [30] .
Development
Oviparous sponges remove zygotes into the external environment, where their development takes place. Many sponges are characterized by live births, in which zygotes remain inside the mother's body and leave it with already formed larvae . In viviparous sponges, embryos develop in special brood chambers formed by flattened cells of choanocytic or amoebocytic origin, as well as endopinacocytes. At the end of development, the walls of the brood chamber take part in the formation of the channels of the aquifer system [104] . Embryos and sponge larvae are lecithotrophic, that is, they feed on yolk [105] . Sponges demonstrate a wide variety of variants of embryogenesis and larval structure, but their development, despite external differences, is very similar at the molecular level (patterns of gene expression, cell movement and death, etc.) [19] .
Crushing
The zygote of the sponges undergoes complete and uniform crushing, however, the relative position of the blastomeres varies among different representatives [106] . In this regard, four main crushing methods are distinguished in sponges:
- disordered (chaotic), characteristic of Homoscleromorpha and most viviparous ordinary sponges;
- crushing with features of radiality is described in egg-laying ordinary sponges, as well as glass sponges;
- polyaxial, described in Halisarcidae (ordinary sponges) and Calcinea (calcareous sponges);
- incurvation crushing is inherent in Calcaronea (calcareous sponges) [107] [108] .
Due to the weak specialization of the egg cell in all sponges (except Calcaronea), crushing, as a rule, is not characterized by the stability of the spatial arrangement of the blastomeres . The synchronism of divisions is either completely absent or is lost early. In the early stages of embryogenesis in many sponges, endosymbiotic bacteria penetrate the embryo from the mesochil. In some cases, somatic cells of maternal origin invade the embryo [109] .
In egg-laying ordinary sponges of the genera Tetilla and Stylocordyla, the development is direct: after crushing, the stage of cell differentiation and the development of a young sponge immediately begin [109] .
Embryonic morphogenesis
During the formation of multicellular embryos (larvae) in sponges, almost all types of cellular movements characteristic of eumetases are described : morula delamination , cell delamination, invagination, and multipolar immigration. However, sponges also exhibit unique morphogenesis not found in other multicellular animals — for example, excursion, the formation of a pseudoblastula due to the introduction of maternal cells into a crushing embryo, polarized delamination, unipolar proliferation, and multipolar emigration [110] .
The embryonic development of the sponges is fundamentally different from that of eumetazoids. During the embryogenesis of sponges, the formation of germ layers does not occur, that is, gastrulation and gastrula are absent, although intensive fragmentation and cell differentiation occurs after crushing [108] . In the late stages of crushing, which is usually called a blastula or morula, differentiation of larval cells begins in sponges. The fate of specialized larval cells of the sponges (integumentary, skeletogenic, and others) is not homologous to the fate of the germ layers of the eumetazoids, because the sponges do not have parts of the body, organs, and tissues homologous to those of the eumetazoids. Thus, sponges do not and cannot have ectoderm and endoderm [111] .
Larvae and metamorphosis
Eight types of larvae have been described in sponges: celoblastula, calciblastula, zincctoblastula, amphiblastula, dysferula, hoplitomella, parenchyma and trichimella [112] .
The simplest larvae are coeloblastula and calciblastula, which have a single-row flagellum and an extensive internal cavity (blastocele) filled with structureless fluid, individual somatic cells of maternal origin and symbiotic bacteria. While the coeloblastula leads a planktonic lifestyle, some flagellate cells lose their flagella and migrate to the blastocele, so that in the end, the hollow coeloblastula turns into a sterroblastula devoid of a cavity [113] . Coeloblastula is described in some ovipositing ordinary sponges, and calciblastula is inherent in calcareous sponges of the subclass Calcinea. In calciblastula, the outer layer of cells may contain individual flagellate cells [112] .
Another single-layer larva is an amphiblastula; it also represents a hollow ball. The anterior hemisphere of the amphiblastula is formed by small flagellum cells, and the posterior one by the flagellum-free large cells with granular cytoplasm. Four cells of the “cross” are located in the equatorial region. In the small internal cavity are various maternal cells and symbiotic bacteria. When the amphiblastula is located in the mesochil of the parent sponge, all its flagella are turned into a blastocele, however, an opening soon forms, through which the larva turns the flagella out (inversion). After that, the larva goes out. Inversion occurs in those sponges in which eggs were formed from choanocytes [113] . Amphiblastules are characteristic of calcareous sponges of the subclass Calcaronea [112] .
Dysferula is one of the larvae of the group of common Halisarca sponges. Inside the larva there is a closed layer of flagellar cells, which was formed as a result of intussusception (invagination) of the superficial flagellum [114] .
Hoplitomella is a unique planktonic larva of the drilling sponges Alectona and . It is devoid of integumentary flagellum cells. Inside the hoplitomella there are choanocytic chambers, various cells, collagen fibers and spicules such as disco trienes, which are absent in adult sponges [115] .
The most complex parenchymal larvae are arranged; their organization can vary even within the same genus. Parenchymal larvae are characteristic of the vast majority of common sponges. The parenchymal is externally covered with a single or multi-row layer of single-flagged cells; in structure it corresponds to the hypothetical phagocytella of I. I. Mechnikov [116] . Inside the larva are sclerocytes, archaeocytes, secretory cells, collenocytes, as well as the larval skeleton, if it is characteristic of an adult sponge. In this regard, the parenchymal can be considered to some extent “almost ready” as a young sponge [113] . Parenchymals entering the external environment are characterized by different degrees of differentiation of larval cells, and sometimes definitive (that is, inherent in an adult sponge) [115] .
Trichimella is characteristic of many glass sponges [113] . The fundamental differences between trichimella and parenchyma are the presence of trichimella in the middle part of the belt of cells carrying up to 50 cilia . The entire larva, including ciliary cells, is covered with a syncytial layer of pinacoderm, with the cilia coming out through openings in the syncytium. Inside the trichymella there are flagellar chambers formed by nuclear-free collar bodies, which differ from those in the adult sponge, since they are not connected by a common cytoplasm. Trichymella has stauroactin spicules absent in adult sponges [117] .
Homoscleromorpha is characterized by a larva of a cytoblastula, which is similar in appearance to a coblastula, since it is formed by a single layer of ciliary cells and has an extensive internal cavity. However, the cells of the outer layer are connected by desmosomes in the apical part, and from the side of the cavity they are underlain by the basal plate, therefore they can be considered as epithelial cells. In the region of the posterior pole of the larva, there is a belt of cells with intranuclear paracrystalline bodies. The inner cavity of the larva contains maternal cells and symbiotic bacteria [118] .
Anteroposterior polarity is characteristic of larvae of all types except hoplitomella. It manifests itself in the shape of the body, the nature of the ciliary cover at different poles, in the uneven color, as well as on the histological level (the nature of the distribution of various types of cells, the orientation and location of spicules in the case of parenchymals, the uneven distribution of symbiotic bacteria in the larval cavity in the case of zincoblastula). Many larval structures die during larval development and are not preserved in adults. Among these structures is the larval ciliary cover, which performs locomotor, attachment, and possibly trophic functions, as well as the larval skeleton [118] .
All sponge larvae are lecithotrophic: they use yolk granules and lipid granules not consumed during embryogenesis as an energy source. Due to lecithotrophy, sponge larvae live relatively shortly [113] . However, the parenchymals of some ordinary sponges can phagocytize organic particles from the external environment using flagellate cells [118] . Usually, the emergence of larvae is stimulated by light and occurs, as a rule, at dawn [113] . Many larvae corkscrew in water. The features of the movement of the sponges are most likely due to the arrangement of the cilia, which varies greatly among different groups [92] .
In the larva of the common sponge there are 3 types of cells that can perform a sensory function. At the posterior end of the larva there are specialized photoreceptor cells that form a ring of pigmented cells with photosensory membranes and long cilia ; they regulate phototaxis . The second type of sensory cells is globular cells filled with vesicles ; they express a number of postsynaptic genes. Finally, at the anterior end of the larva there is a group of cells equipped with one cilium. They are connected to other cells by special contacts, but no chemical synapses were detected [72] .
Sponge larvae can quickly respond to environmental stimuli through a series of orientational behavioral programs such as geotaxis , phototaxis and rheotaxis . In addition, they can choose a substrate, which subsequently sink [72] .
After a certain time (from several hours to several days), the larvae settle to the bottom and begin to search for a suitable place for attachment [113] . The metamorphosis of sponge larvae begins with the attachment of the larva to the substrate. Next, exopinacoderm and basopinacoderm are formed, choanocytic chambers (or choanoderm in asconoid representatives) develop and channels connecting with them form pores and an oskulum. At the same time, synthesis of sponging, spicules and the main substance of mesochil occurs. The single-spit individual, which forms directly from the larva and has an aquifer system different from that in adult forms, has an asconoid structure in calcareous sponges and is called olintus. Olintus has a single saccular choanocytic chamber and apical oskulum. In ordinary sponges and Homoscleromorpha, this stage has a siconoid or leukoid structure and is called ragon; it has one oskulum and several choanocytic chambers [108] . In most parenchymal larvae, metamorphosis results in the destruction of integumentary cells, and new integuments of the young sponge are formed by internal cells. In amphiblastula, calciblastula, coblastula, zincctoblastula and Halisacra larvae, only external cells differentiate during larval morphogenesis [119] .
Types of Development
Under the type of development is meant a complex of inherited and acquired morphogenetic processes connected with each other. In sponges, 8 types of development are known, named for characteristic larvae. They are listed in the following table.
| Type of | Taxon | Egg | Splitting up | Larva | Metamorphosis | Asexual reproduction | Source |
|---|---|---|---|---|---|---|---|
| Amphiblast | Calcaronea (calcareous sponge) | Isolecital, oligolecital, oval | Complete, Asynchronous, Uneven, Incursive | Amphiblastula | According to the mesenchymal type: flagellum cells plunge inside and give rise to choanocytes, mesokhil cells, sclerocytes and endopinacocytes, granular flagellate cells give exo- and basipinacoderm | Budding | [120] |
| Calciblast | Calcinea (calcareous sponge) | Isolecital, oligolecytal, without signs of polarization, no special feeding cells | Full, uniform, polyaxial, there is a crushing cavity | Initially, the whole-blast larva acquires an anteroposterior polarity and becomes a calciblastula. The main type of larval cells is flagellum cells. There are special intercellular contacts. | According to the mesenchymal type | Budding | [121] |
| Trichimelial | Glass sponges | Isolecital, oligolecytal, without signs of polarization, no special feeding cells | Full, uniform, asynchronous | Trichimella, there are cork contacts | Not described | Different types | [122] |
| Dysferal | Halisarcida (ordinary sponge) | Isolecital, multilecital, without signs of polarization, no special feeding cells | Complete, uniform, asynchronous, polyaxial, there is a crushing cavity. | Celoblastula, parenchyma, dysferula. Larvae are completely covered with flagella. Adhesive contacts available | Mixed: exopinacoderm develops according to the epithelial type from flagellar cells of the posterior pole of the larva, and basopinacoderm, ecdynacoderm and choanoderm along the mesenchymal | Not known | [123] |
| Parenchymal | Ordinary sponges | The structure varies | Complete, asynchronous, disordered | Parenchyma | According to the mesenchymal type, it is accompanied by the migration of flagellum cells inward and outward. Full or partial phagocytosis of flagellar cells of the larva | Budding, fragmentation, hemmulogenesis | [124] |
| Direct development | Ordinary sponges | Small, isolecital, oligolecital, without signs of polarization | Full, uniform, asynchronous | Morula delamination | No | Budding | [125] |
| Whole-blast | Ordinary sponges | Eggs are surrounded by a layer of collagen and mucus, isolecital, oligolecital, non-polarized | Complete, uniform, may be radial | Coeloblastula; adhesive intercellular contacts | A part of flagellum cells is evicted inward and forms choanocytes and pinacocytes that remain on the surface - pinacocytes | Budding | [126] |
| Zincoblast | Homoscleromorpha | Isolecital, multilecital, no polarity, no specialized feeding cells | Complete, uniform, asynchronous, disordered | Initially, a celoblastula that turns into a zincoblastula. Basal plate, desmosomes | Epithelial type | Budding and fragmentation | [127] |
Amphiblastular type of development. 1 - egg, 2 - incurvation crushing, 3 - stomoblastula, 4 - incurvation, 5 - amphiblastula, 6 - olintus
Calciblastular type of development. 1 - egg, 2, 3 - polyaxial crushing, 4 - coeloblastula, 5 - calciblastula, 6 - the beginning of metamorphosis, 7 - olintus
Trichimelial type of development. 1 - egg, 2, 3 - radial fragmentation, 4 - blastula, 5 - delamination, 6 - fusion of macromers with the formation of trabecular syncytium, 7 - differentiation of cell types, 8 - longitudinal section of the larva, 9 - appearance of trichimella
Direct development. 1 - zygote, 2 - radial fragmentation, 3 - formation of exopinacoderm, 4 - young sponge
Dysferal type of development. 1 - egg, 2, 3 - polyaxial crushing, 4 - intussusception, 5 - dysferula, 6 - ragon
One of the options for parenchymal development. 1 - egg, 2 - chaotic fragmentation, 3 - morula, 4 - parenchymal, 5 - metamorphosis
Holoblast type of development. 1 - egg, 2 - crushing, 3 - morula, 4, 5 - coeloblastula, 6 - metamorphosis
Zincoblastular type of development. 1 - egg, 2 - crushing, 3 - morula, 4 - coeloblastula, 5 - zinctoblastula, 6 - metamorphosis
Life
The lifespan of sponges varies in different species from several weeks and months to tens and hundreds of years. In temperate latitudes, it ranges from one year to several years, and freshwater sponge species usually live only a few months, but in some cases they can create perennial formations of a special kind. However, tropical marine sponges, as well as deep-sea species, live very long - up to 200 years or more [26] [128] . The long-livers among the sponges are probably - the specimen obtained in 1986 in the East China Sea at a depth of 1110 m is estimated to be 11 (± 3) thousand years [129] .
The ability to reproduce sexually, some species acquire at the age of 2-3 weeks, and others only when they reach the age of several years. In general, sponges grow relatively slowly (the highest growth rate was observed in forms with a short lifespan). The growth and age of the sponges also greatly depends on external factors - for example, a one-year temperature difference. Some sponges (Sclerospongia) have a very low growth rate, giving an increase of about 0.2 mm per year, so that the age of such specimens with a diameter of 1 m can be 5000 years [26] [128] . At the same time, some species grow at a rate of several decimeters per year. A young sponge grows relatively quickly, but with an increase in body volume, its growth rate decreases [130] .
Molecular Biology
The features of the sponge genomes were studied by the example of ordinary sponges and . They have relatively large genomes (1670 megabases, the mass of genomic DNA is about 1.7 pg ). In addition, it was shown that the genomes of sponges consist mainly of sequences represented in the genome only once (without repeats) [131] . Sponge genes contain few introns , especially for genes encoding proteins , and their introns are quite short: most of them are 0.1–2 kilobases long, and none of the introns found has a length of more than 5 kb [132] . The nuclear genome of sponges contains the genes that make up homeobox genes. The situation with the presence in sponges of other key for multicellular animals homeobox genes — Hox genes — is less clear [133] . In many sponges, the chromosomes are very small, and microchromosomes are abundantly represented in karyotypes [20] .
There are introns in mitochondrial DNA (mtDNA) of sponges, although most Metazoa do not have introns in mtDNA (exceptions are only corals and sea anemones) [134] . In calcareous sponges, mtDNA has a linear structure, rather than a ring one, as in most Metazoa. So, in mitochondrial DNA is represented by 6 linear chromosomes. In addition, tRNA modification occurs in this sponge in mitochondria, and rRNAs are fragmented [135] . It should be emphasized that different sponges in mitochondria show different differences from the standard genetic code . So, in glass sponges, the AGR codon does not encode arginine , but serine , and in the calcareous sponges , Sycon coactum and Petrobiona massiliana - glycine or serine. In addition, in P. massiliana, the AUA codon does not encode isoleucine , but methionine . In the calcareous sponges Clathrina clathrus and the UAG codon does not terminate , but encodes tyrosine , and the CGN codons (N is any nucleotide ) encode glycine instead of arginine . Finally, in the calcareous sponge Leucosolenia complicata, AGR codons began to encode methionine [136] .
Sponges have identified many genes and proteins responsible for organizing their body. Adhesion molecules have been identified that ensure the interaction of cells with each other, as well as with the intercellular matrix . The most important molecule that provides the interaction of two cells is the aggregation factor (AF). provides an association of AF with a membrane cell aggregation receptor (AR). The key proteins of the intercellular matrix - collagen - and fibronectin-like proteins - interact with integrins located in the cell membrane . At least one morphogen has been identified in sponges - ; most likely, it is involved in the formation of the body axis. In addition, many protein families characteristic of other Metazoa appear in sponges: intercellular matrix proteins (collagen, proteins containing FN 3 -, SRCR, SCR domains ); proteins involved in signal transmission ( tyrosine kinases ); innate immunity proteins ( cytokines , (2-5) A synthetase, variable domains) [21] . Thus, molecular biology data indicate that sponges are not unorganized clusters of cells, but complexly organized animals with a specific body plan [137] .
Sponges do not have real neurons and the nervous system , but they have genes associated with the functioning of neurons, behavioral responses to external stimuli and sensory cells involved in signal perception and expressing genes characteristic of neurons. common sponge , as well as analysis of transcriptomes of sponges of various classes, has shown that sponges have many genes that are characteristic of cells of the striking and bilaterally symmetrical animals of the nervous system. Among them are genes associated with fast synaptic transmission: potential-dependent ion channels , neurosecretory proteins of the SNARE group (syntaxin1, SNAP-25 , synaptobrevins ); genes associated with the primary determination of neurogenic fields (including proteins); enzymes associated with the synthesis of neurotransmitters (including aromatic amino acid decarboxylase and tryptophan hydroxylase ); most of the genes of postsynaptic endings ( NOS , citron and ) [72] . In this case, the sodium channels available in choanoflagellates are absent in sponges [36] .
Distribution
Sponges are spread all over the world. Most species inhabit ocean waters from the polar regions to tropical regions [26] [86] . At the same time, sponges reach the greatest species diversity precisely in tropical and subtropical regions of the World Ocean [26] . 219 [138] species (according to other sources, about 150 [28] ) live in fresh water bodies.
Most species live in calm and clean waters, since particles of silt and sand suspended in the water or raised from the bottom by currents can clog pores in the bodies of the sponges, thereby hindering the processes of respiration and nutrition [139] . Since the sponges lead an attached sedentary lifestyle, they need a solid substrate for their development and growth. In this regard, clusters of sponges occur in places where rocky materials are present at the bottom: stones, boulders, pebbles, etc. [26] However, some species can be fixed in soft sedimentary bottom sediments with the help of a root-like base of the body [140] [28 ] ] .
Most sponges live at shallow depths: up to 100-500 m. With an increase in depth, the number of sponge species decreases. At depths greater than 1000-1500 m, sponges are usually quite rare, so the number of deep-sea species is small [26] [28] .
In temperate waters, sponges are more numerous, but less diverse than in the tropics [26] . This is probably due to the fact that many organisms that feed on sponges are found in the tropics [141] . Glass sponges are most abundant in polar waters, as well as at great depths of temperate and tropical seas, because their porous body structure allows them to extract food particles from these food-poor waters with minimal cost. Ordinary sponges and calcareous sponges are numerous and diverse in calmer non-polar waters [27] .
The sponges that live in the tidal zones are well adapted for a short stay in the open air when they emerge from the water at low tide. At the same time, their mouths and pores are closed, which prevents excessive loss of moisture and drying out [26] .
Among the endemic species, the sponges of Lake Baikal [142] , represented by the endemic family , which unites 4 genera and 13 species [143], stand out.
Differences in habitats of different classes of sponges are indicated in the following table [86] :
| Type of water [86] | Depth [86] | Type of surface [86] | |
|---|---|---|---|
| Lime sponges | Marine | Less than 100 m | Solid |
| Glass sponges | Marine | Great depths | Soft or hard precipitation |
| Ordinary sponges | Sea, fresh (about 80-90% of all known types of sponges) [144] | From the intertidal zone to the abyssal ; [86] predatory common sponges can be found at a depth of 8840 m [81] | Any |
Sponge Reefs and Communities
Sponge reefs are a special type of reef [145] . Similar reefs were previously widespread in the history of the Earth [146] . The earliest of them date from the Middle Triassic (245–208 million years ago), and they reached their greatest prosperity at the end of the Jurassic period (208–146 million years ago). At this time, the sponge reef system, formed mainly by six-pointed sponges and having a length of about 7 thousand km, crossed across the northern basin of the ancient Tethys Ocean ; it is the largest among the known designs ever created by living organisms. A sharp decrease in the number of sponge reefs occurred in the Cretaceous against the backdrop of the ever-increasing dominance of coral reefs [147] .
Shallow-water sponge reefs are now absent, and all known similar reefs are deep-sea [148] . Modern sponge reefs need unique living conditions that can explain their rarity. They arise in the trenches of the continental shelf , in places with optimal conditions for their development: with a relatively low sedimentation rate, with high flow rates [146] , a high concentration of silicon dioxide in water and a low concentration of oxygen, with a stable seabed consisting of stones, coarse gravel and large boulders, with a water temperature of 5.5-7.3 ° C [147] . Sponge reefs are currently known in the west of the Canadian continental shelf [149] , are widespread in the North Atlantic, in the Canadian Arctic and on the continental shelf in the Antarctic [148] . Formed by silicon sponges ( Demospongiae ), the reef exists in the Arctic Ocean off Axel Heiberg Island [150] . Four reefs formed by six-pointed sponges in Queen Charlotte Bay at a distance of 70–80 km from the coastline and at a depth of 165–240 m occupy an area of 700 km² of seabed [148] . Some sponges from the genus in the North Pacific Ocean can create reefs over 20 meters high, extending several kilometers. Also along the deep-sea coral reefs , a zone of coral fragments usually stretches, in which sponges dominate, among which there are species that destroy dead corals [151] . In the southwestern part of the Barents Sea, there are deep-sea communities of sponges and brachiopods [146] .
Relationships with Other Organisms
Symbiotic Associations
Sponges can enter into symbiotic relationships with representatives of most groups of organisms [152] : with algae [153] , mushrooms [154] , prokaryotes [155] , etc.
Representatives of the Porifera type provide habitat for many unicellular and multicellular organisms. Multicellular organisms usually act as commensals and settle on the surface of the body of the sponge and in its internal cavities, without interfering with its vital functions [156] . Among the endobionts of sponges, scarabidae , turbellaria , nemerthins , sipunculides , polychaetes , mollusks , crustaceans , multi- legged , echinoderms , ascids, and fish were noted [157] .
Dinoflagellates , zoochlorella , red algae , filamentous green algae , and also mushrooms can be unicellular symbionts of sponges [156] . Freshwater sponges are most often found in symbiotic relationships with various algae, which determine their green color [26] .
The sponges in which photosynthetic endosymbionts settle, emit three times more oxygen than they absorb, and also produce more organic matter than they consume. This role of sponges in the formation of primary production is especially important in the Australian Great Barrier Reef , but in the Caribbean it is not so great [158] .
However, especially important symbionts of the sponges are obligate species-specific autotrophic or nonautotrophic prokaryotes (bacteria and archaea ) that inhabit the mesochilus. The interaction between the sponges and these microorganisms can occur in different ways. The uniqueness of the symbiosis of sponges with prokaryotes lies in the fact that all the sponge species studied so far have symbiotic associations with one or more types of bacterial symbionts [155] [159] [160] [161] .
Prokaryotic symbionts, as a rule, are located between the elements of the mesochil and are physically separated from the water by the pinacoderm. However, sponges in which symbionts are intracellular are known: they are localized in special cells - bacteriocytes [162] . The biomass of prokaryotic endosymbionts can be up to 40% (and even up to 60%) of the mass of the host sponge, and the concentration of bacterial symbionts in the sponge can be 2-3 orders of magnitude higher than the concentration of prokaryotic organisms in the surrounding aquatic environment [163] . Prokaryotes can penetrate the body of the sponge in two ways: from water in the environment, or by vertical transfer from the mother sponge to the next generation through eggs or larvae; the possibility of vertical transfer of prokaryotic symbionts indicates their exceptional importance for the life of sponges. Sponges manage to maintain diverse, but at the same time specific microbial communities within themselves, despite the fact that many marine microorganisms penetrate their bodies along with the flow of water during filtration nutrition [164] .
Inside the sponges, prokaryotes of the most diverse systematic position, belonging to about 20 bacterial fila, settle, and representatives of one phylum - Poribacteria - are found exclusively in sponges. Symbiotic prokaryotes can take part in various physiological processes of sponges - for example, in the cycle of insoluble proteins and structural rearrangements of the organic skeleton and extracellular matrix. Many symbiotic prokaryotes synthesize antibiotics and pungent odor repellent substances. They can also serve as an additional source of nutrition for sponges, moreover, they can be absorbed both by prokaryotes by phagocytosis and their metabolic products [165] . We can say that the host sponge and microorganisms inhabiting it are metabolically coupled and exchange nitrogen compounds, CO 2 fixation products, secondary metabolites, and organic compounds obtained from water [164] . For example, inside the sponge Lophophysema eversa autotrophic microorganisms live, which are able to oxidize ammonium , nitrite and sulfide and therefore play an important role in the circulation of carbon, nitrogen and sulfur in the body of the sponge. In addition, they neutralize ammonia toxic to the host sponge, nitrite, and sulfide [166] .
Sponges come in a variety of relationships with other animals. For example, a fairly large sponge , living on stony substrates, can also inhabit thickets of marine grasses . At the same time, it allows you to surround and grow yourself in sponges that live in sea herbs and are dangerous for local starfish ; Thus, Lissodendoryx colombiensis provides protection for itself, and the sponges of thickets of sea grasses are able to live at some elevation relative to the seabed [167] .
The relationships between crustaceans and sponges are especially diverse [26] . Many species of small copepods and isopods , amphipods cohabit or parasitize in sponges. Often decapods and shrimps use sponges as a refuge from enemies. Shrimp of the genus Synalpheus from the Alpheidae family form colonies inside the sponges, with each shrimp species inhabiting only a certain type of sponge; Due to this, the genus Synalpheus is one of the most numerous genera of crustaceans . Species Synalpheus regalis uses sponges not only for nutrition, but also for protection against other shrimp and predators. 16,000 shrimp individuals can live in one sponge, feeding on the largest particles that enter the sponge’s body during filtration feeding [168] [169] . In another shrimp family, Spongicolidae , almost all representatives are obligate symbionts of deep-sea glass sponges (only the genus free-living forms) [170] . In particular, the male and female from the genus Spongicola settle in the cavity of the glass sponge Euplectella oweni at the larval stage, and in the adult state they can no longer leave the sponge skeleton [26] .
Unattractive sponges for predators are often used by some crabs (for example, Dromia personata ), which carry pieces of sponges on themselves, supporting them with the back pair of their limbs. Over time, the sponge takes the form of a shell [171] [172] . The sharp-pointed crab attaches sponges, algae, and sedentary animals to its carapace, so that a micro-community masking the crab forms on its back [173] . 13 species of hermit crabs can form a symbiosis in the form of commensalism with a cork sponge ( Suberites domuncula ) [174] . The latter settles on the empty shell of the gastropod mollusk, and the hermit crab settles in the same shell. Over time, the entire shell is overgrown with a sponge; inside the latter, a spiral cavity forms in which the hermit crab lives [26] .
Some animals use separate sponge needles as protection, which are deposited in bulk after their death at the bottom. These animals build their houses partially or completely from spicules and sponge needles ( foraminifera , some worms, holothurium Pseudostichopus trachus , etc.) [26] .
Sometimes sedentary, attached animals, which include various barnacle cancers (in particular, balyanus ), can settle on the lips. Sea ducks are often found on the root needles of a number of glass sponge species, and colonial corals (Palythoa) can also settle. Cortical sea anemone ( Parazoanthus axinellae ) is constantly settling on some sponges ( Axinella ) [26] .
Natural enemies and defenses against them
Many sponges have spicules that protect them from being eaten by echinoderms and other predators [158] . However, sea turtles eagerly feed on sponges, and silica spicules can make up up to 95% of their feces. Sponges are typical passive-poisonous animals that use their toxic metabolites to protect themselves from enemies [176] , including natural haloalkaloids [177] . The first toxin isolated from sponges ( ) was a substance called suberitin [176] . The toxins of the bromofackelin group were first isolated from the sponge ; a toxin close in structure to dibromisophakelin was found in the sponge [177] . Pyrrole alkaloids are widely found among sponges, especially representatives of the family [177] . It was shown that 9 out of 16 Antarctic sponges and 27 out of 36 Caribbean sponges are toxic to fish . At the same time, species specializing in feeding on sponges are found among fish ( Moorish idol , angel fish of the genus [178] , butterfly fish of the genus [179] , etc.) [86] . The nudibranch mollusks of the exhibit a narrow specialization, eating only certain types of sponges.
Toxins secreted by the sponges can also be directed against other sedentary organisms (for example, bryozoans and ascidia ) so that they do not settle on the sponges and in the immediate vicinity of them, so that the sponges can successfully conquer new areas for life. and dibromagelaspongin synthesized by sponges from the genus examples of such toxins. The Caribbean sponge emits toxins that kill coral polyps , and the sponge grows on their skeletons. Organic compounds similar in structure to hymenialdisines , as well as 2-bromaldizine, characterized by a moderate antimicrobial effect, were found in various types of sponges [177] .
Many sponges have a strong odor, for example, the “garlic” sponge . Some species can cause dermatitis in humans through direct contact [176] . For example, touching the Caribbean “fiery” sponge can cause severe skin irritation [180] .
Glass sponges do not synthesize toxins, because there are few predators at great depths in which they live [139] .
There is a family of winged insects - sisyridae (Sisyridae), representatives of which parasitize on freshwater sponges. The female lays eggs on plants hanging over the water. After hatching, the larvae fall into the water and look for sponges that they will feed on. Their elongated oral apparatus is designed to penetrate the sponge and suck fluid from her body. In some species, the larvae attach to the surface of the sponge, while in others they settle in the internal cavities of its body. A fully formed larva emerges from the water and twists a cocoon in which it pupates [181] .
Abdominal mollusks from the pleurotomarium family feed mainly on sponges [182] . Some starfish also specialize in sponge feeding (for example, species of the genus ) [183] .
Diseases
| External Images | |
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| Aplysina Sponge Red Strip Syndrome | |
Most often, diseases are noted among the sponges of Papua New Guinea , Mexico and the Great Barrier Reef . Despite the enormous environmental and economic importance, sponge diseases are poorly understood. It is known that fungi , viruses , and bacteria (including strains of the genera Bacillus and ) can act as pathogens . An association of disease epidemics among sponges and environmental conditions, such as climate change, has been shown [184] .
Caribbean sponges of the genus suffer from a particular disease known as red streak syndrome. On the bodies of diseased sponges, one or several strips of rusty color are formed, sometimes surrounded by strips of dead tissue. The body of the sponge can be completely surrounded by such lesions. The disease is apparently contagious, and about 10% of Aplysina cauliformis individuals are infected on the Bahamas reefs [185] . Strips have a bright color due to cyanobacteria , but it is not known whether it is responsible for the occurrence of the disease [185] [186] .
Other
The drilling sponges of the family secrete special substances produced by archaeocytes and allowing them to dig into stones, corals and shells of dead mollusks [187] .
Bottlenose dolphins from Shark Bay in Western Australia use sponges in an interesting way: they attach sponges to their rostrum to protect it from damage while searching for food in the sandy bottom [188] . This type of behavior, called sponging, is known only among bottlenose dolphins from Shark Bay, and more often than not sponging is demonstrated by females. It has been shown that mothers teach sponging to their daughters, with all dolphins showing sponging being close relatives to each other (so sponging seems to be a recent acquisition) [189] [190] .
Evolutionary History
Fossil Residues
It is known that ordinary sponges, but not eumethasoys, can synthesize , a stable derivative of 24-isopropyl cholesterol . It has been shown that the synthesis of 24-isopropylcholestan is also incapable of choanoflagellates, the closest unicellular relatives of animals, therefore, the presence of 24-isopropylcholestane in fossil residues can be used to judge the existence of sponges at that time [191] . According to 2005 data, the oldest findings of 24-isopropylcholestane are 1800 million years old [192] , but a more accurate analysis showed that this biomarker appeared only shortly before the end of the Proterozoic glaciation , that is, 635 million years ago [193] . Thus, there is no evidence of sponges before the Proterozoic glaciation. Moreover, 24-isopropylcholestane, the “sponge biomarker,” could also be synthesized by other organisms, for example, seaweed [194] .
Although molecular clocks and biomarkers indicate that sponges existed long before the Cambrian explosion , silicon spicules, similar to ordinary sponge spicules, appear in the geological record only from the Cambrian [195] . However, there are unconfirmed data on the finds of silicon spicules in stones having an age of 750 million years [196] . The well-preserved ancient fossil remains of the sponges found in the Doushanto formation are 580 million years old and belong to the Ediac period . These fossil remains, in which spicules, pinacocytes, porocytes, archeocytes, sclerocytes and internal cavities are distinguishable, were assigned to ordinary sponges. Other possible ordinary sponges were found in Early Cambrian deposits in the Maotian Shan shales and were 525–520 Ma old [197] . The oldest fossils of glass sponges are 540 million years old and were found in rocky rocks in Australia, China and Mongolia [198] . In the early Cambrian sponges from Mexico belonging to the genus Kiwetinokia , the fusion of several small spicules to form one large spicule was confirmed [199] . Calcium carbonate spicules inherent in calcareous sponges were found in Australia in the early Cambrian rocks from 530 to 523 Ma. Freshwater sponges appeared much later. The earliest fossils of freshwater sponges belong to the Middle Eocene and are from 48 to 40 Ma [198] . Although 90% of modern sponges belong to ordinary sponges, fossil remains of representatives of this group are rarer than remnants of sponges of other groups, since their skeleton is formed by relatively soft, poorly fossilized spongin [200] . The appearance of endosymbionts inside the sponges belongs to the early Silurian [201] .
Phylogeny
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As already noted in the “Study History” section, for most of the 20th century sponges (Porifera type) were included in Metazoa (real animals) and contrasted as all other representatives of Parazoa group ( Eumetazoa ) [16] . At the same time, Porifera monophilia was usually not called into question [203] , but with regard to Metazoa as a whole, such doubts arose. In 1970-1980 a number of zoologists adhered to the hypothesis of the independent origin of sponges (together with choanoflagellates ) and Eumetazoa from unicellular ancestors (presumably from ) [204] [205] .
The use of molecular phylogenetics has put an end to doubts about the unity of Metazoa; it was also revealed that the choanoflagellates are the sister group for Metazoa as a whole [206] . However, the data of molecular studies regarding monophilia of sponges were contradictory. Thus, the results of a number of studies obtained since 2001 have shown that some groups of sponges are systematically closer to eumetases than other groups. This meant Porifera paraphilia : it turned out that the last common ancestor of the sponges was the ancestor of not only the sponges, but also the eumetazoids. An analysis of rRNA genes carried out in 2001 showed that the most fundamental differences inside the sponges are between glass sponges and all other sponges, and eumetazoids are closest to calcareous sponges [203] . Two analyzes of 2007, based on differences in rRNA genes and in spicules, respectively, showed that glass sponges and ordinary sponges are much closer to each other than each of these groups to calcareous sponges, and the latter, in turn, are closest to eumetazoyam [198] [207] .
Other anatomical and biochemical data indicate the proximity of eumetasoids and Homoscleromorpha, a class of sponges that has long been considered as part of ordinary sponges. An analysis of nuclear DNA in 2007, which did not include glass sponges and ctenophores , showed that the closest group of sponges to eumetases is the Homoscleromorpha class, the next closest are calcareous sponges, and ordinary sponges are evolutionary “aunts” for both groups [208] . Homoscleromorpha spermatozoa have a number of similarities with eumetazoic spermatozoa, which are absent in all other sponges. In both Homoscleromorpha and eumetazoids, cell layers attach to a carpet-like basement membrane, consisting mainly of type IV collagen protein, which was not found in the remaining sponges. However, spongin, which supports the common sponge mesocile, is close to type IV collagen [209] .
So, according to the above data, sponges are the earliest branches of the clade of true multicellular animals (Metazoa). However, in 2008, 150 genes of 21 species were analyzed (from mushrooms to humans; the sponges in this analysis were represented by only two species), which showed that at the very base of the Metazoa tree are not sponges, but ctenophores. If his data are correct, then either the ctenophores obtained all their complex structures independently of Metazoa, or the ancestors of the sponges had a much more complex structure, and all modern sponges are radically simplified forms [210] . The data obtained later support the version that ctenophores are a sister group to the other metazoans [211] [212] [213] [214] . Последние данные по морфологии, особенностям развития и молекулярной биологии (в частности, анализ митохондриальной рибосомной ДНК, а также 18S и 28S ядерной митохондриальной ДНК [215] ) говорят о том, что губки всё же являются монофилетической группой и вместе со стрекающими выступают как сестринские группы по отношению к двустороне-симметричным животным [216] .
В пределах типа губок стеклянные и обыкновенные губки считаются наиболее близкими друг к другу как по молекулярным, так и по анатомическим и палеонтологическим данным. Подтверждена монофилия стеклянных губок, известковых губок и двух подклассов известковых губок (Calcinea и Calcaronea). Однако филогенетические отношения в пределах обыкновенных губок подверглись существенным изменениям. Чётко монофилетическая группа Homoscleromorpha была выделена из обыкновенных губок в ранге класса, а оставшиеся в составе Demospongiae отряды и семейства также образуют монофилетическую группу, причём они подразделяются на четыре кластера уровнем выше отряда , рассматриваемые сейчас в ранге подклассов [217] . Что же касается типа Porifera в целом, то имеющиеся данные не позволяют сделать однозначный вывод, является ли данный таксон монофилетическим или парафилетическим ; в тех исследованиях, которые свидетельствуют в пользу монофилии губок, Homoscleromorpha выступают как сестринская группа по отношению к известковым губкам. В любом случае все четыре выделяемых ныне класса губок дивергировали друг от друга свыше 600 млн лет тому назад [218] .
Археоциаты — это группа животных, чьи ископаемые остатки весьма многочисленны в отложениях раннего кембрия (от 530 до 520 миллионов лет назад), но полностью отсутствуют в отложениях позднего кембрия. Им приписывали родство с губками, стрекающими, водорослями и фораминиферами и предлагали выделить в отдельный тип (Archaeocyatha) животных или даже царство (Archaeata или Inferibionta). Однако с 1990-х годов археоциатов рассматривают как своеобразную ископаемую группу губок [219] .
Ранее к губкам относили группу кембрийских мешковидных организмов, известных как . На основании сходства склеритов Chancelloriidae и спонгиновых волокон обыкновенных губок эти две группы сближали [220] , однако в 2002 году появились данные о том, что Chancelloriidae нельзя рассматривать в составе губок, потому что эти организмы являются промежуточным звеном между губками и более высокоорганизованными животными. В качестве одного из доказательств последнего положения приводилось строение покровов Chancelloriidae: они более толстые и плотные, чем у губок [221] . В 2008 году был проведён детальный анализ склеритов Chancelloriidae, который показал, что они очень близки по строению к склеритам — группы подвижных двусторонне-симметричных животных, похожих по виду на звенья кольчуги и известных по ископаемым остаткам из раннего и среднего кембрия [222] .
Classification
Карл Линней, относивший большинство сидячих животных к порядку Zoophyta класса Vermes, ошибочно поместил род обыкновенных губок Spongia к растениям в порядок Algae (водоросли) [223] . Впоследствии долгое время губок выделяли в отдельное подцарство Parazoa царства животных, противопоставляя их эуметазоям, к которым относятся все остальные животные (кроме пластинчатых ) [219] .
В настоящее время известно около 8000 (по другим оценкам, более 15000 [138] [224] ) видов губок, из которых 219 [138] (по другим данным, 150 [32] ) — пресноводные. Современные виды губок разделены на четыре класса [225] .
Класс Известковые губки
В состав класса известковые губки ( лат. Calcarea ) входит 680 видов, объединённых в 5 отрядов ( , , , Clathrinida , Murrayonida [226] [227] ), 24 семейства и 577 родов, что составляет примерно 8,2 % от всех описанных видов губок. Класс признан монофилетическим. Известковые губки — исключительно морские организмы, населяют все акватории от литорали до батиали . Минеральный скелет данных губок состоит только из карбоната кальция. Спикулы (двух-, трёх- и четырёхлучевые) располагаются в мезохиле свободно; иногда спаянные друг с другом спикулы формируют плотный скелет. В классе представлены все варианты строения водоносной системы. Личинка однослойная, с полостью внутри (амфибластула или кальцибластула). Класс подразделяется на два монофилетических подкласса — Calcaronea (личинка амфибластула; первые три перечисленных выше отряда) и Calcinea (личинка кальцибластула; два оставшихся отряда) [225] .
Класс Homoscleromorpha
Sponges of the Homoscleromorpha class used to be classified as ordinary sponges as part of the subclass, however, modern phylogenetic studies show that this is a clearly limited group of sponges, independent of ordinary sponges. This class includes one detachment of Homosclerophorida with two families and 6 genera. Representatives of the class are small smooth sponges with a body shape from cortical to lumpy, living in the seas at shallow depths. The aquifer system is syleibid or leukoid. There may be an inorganic silica skeleton represented by small four-axial spicules and their derivatives, and the rays of the spicules can branch and bend. The basal part of the choanoderm and pinacoderm cells is underlain by a basal plate, the cells themselves are connected by specialized contacts. The basal lamina contains type IV collagen, tenascin and laminin. This structure of the choanoderm and pinacoderm corresponds to the structure of the epithelium of the eumetazoids. During the breeding season, the basal lamina covers the spermatocyst and is present in larvae. Exo- and endopinacocytes are equipped with flagella. A slight variety of cell types is characteristic. The sponges included in the class are viviparous, the larva of the cytoblastula. Asexual reproduction - in the form of budding and fragmentation [228] .
Class Ordinary Sponges
The class of ordinary sponges ( lat. Demospongiae ) includes 83.3% of all modern sponges - about 6900 species (combined in 12 orders: , Astrophorida , , , , , , , , , , [226] [227] ). Sea and freshwater inhabitants. The skeleton is only spongy or spongin with silicon spicules, which are divided into macrosclera and microsclera depending on size. Spongin forms individual fibers or combines spicules. Some forms are completely devoid of skeletal elements. The shape of the body is diverse; the aquifer system is mainly leukoid, but the predatory sponges from the order have completely lost the aquifer system. Larvae are mainly parenchymals, but some groups have single-layer larvae. In different forms, live birth, ovipositor development and external development are found. Four subclasses are distinguished in the composition of Demospongiae: (orders Dendroceratida and Dictyoceratida), Myxospongiae (or : orders Chondrosida and Verongida), Haploscleromorpha (order Haplosclerida) and (common) ( 22) ] [230] .
Grade Glass Sponges
The class of glass sponges ( lat. Hexactinellida, Hyalospongia ) includes about 600 species (7.3% of all the described species of Porifera), collected in 6 orders ( , , , , , [226] [227] ), 26 families and 166 genera. They live in the seas at depths from 5 to 6770 m. Skeletal elements are represented mainly by six-beam spicules and their derivatives. Macroscler fused together form rigid skeletal structures. Most cells combine into syncytium, and individual cells located in pockets or capsules of syncytium can be represented by special pore contacts. The aquifer system is siconoid, silleibid or leukoid. All representatives of the class are viviparous, the larva of Trichimella. There are 2 subclasses of glass sponges: and [231] . The names of the subclasses are associated with the shape of the microscler: in the representatives of the subclass Amphidiscophora (contains the first of the listed orders), the microsclera are amphidisks (thin rods, at the two ends of which there are corollas of several blades), and in the lips of the subclass of Hexasterophora (5 remaining orders) - hexastras ( small six-pointed needles, in which the rays often carry a variety of appendages at the ends) [232] .
Threats and security
A number of factors can adversely affect the size of sponge populations, cause their oppression and death. Currently, the main threat to the reduction in the number of many species is human economic activity. In particular, a trawl fishery is a threat to sponges, in which sponge catches can reach 5-12.5 tons per nautical mile . Under its influence, in the middle of the 20th century, in just a few decades, in many places of the oceans there was a sharp degradation of sponge settlements; Numerous facts are known of the complete destruction of sponge biotopes . As a result of this type of fishing in the Barents Sea, the total biomass of sponges (mainly and ) decreased by two orders of magnitude [233] , and in the central part of the sea mass settlements of sponges almost completely disappeared [234] [235] .
In 2002, a voluntary and regulated prohibition of bottom trawling was adopted in areas where deep-sea sponge reefs are known. However, the voluntary rejection of fishermen is not a fully effective method [236] .
A danger to sponges is also pollution of the world's oceans , especially coastal waters in densely populated regions. Thus, an increase in eutrophication due to pollution by wastewater of the coastal shallow waters of the Black Sea and the subsequent decrease in oxygen concentration in water led to a decrease in the abundance of the Halichondria panicea sponge ( sea loaf ), which is currently listed as a vulnerable species in the Arctic and North Atlantic, in The Red Book of the Black Sea [237] .
Human Use
Calcium carbonate or silica spicules make most sponges too stiff for any use, but two sponges - Hippospongia and toilet sponge ( Spongia ) - have skeletons that are soft. In Europe, these sponges have long been widely used. They were used as a helmet lining, a portable drinking vessel, and also as filters for city water. Before the invention of synthetic sponges, they were used for washing, as a means for applying paint and glaze . In zootechnics and veterinary medicine, Euspongia sponges were used to collect sperm during artificial insemination of livestock [238] . Dry ground sponges served as an abrasive material for grinding metal, wood, and clay products [239] . However, due to overfishing by the mid-20th century, sponges and related industries were on the verge of extinction [240] .
Since ancient times, sponges have been used in medicine. The mildest and thinnest species of the genus Euspongia have been used in surgery to stop bleeding , and also as a contraceptive to prevent pregnancy [238] . Today, sponges retain important medical significance, since they themselves and the endosymbionts inhabiting them synthesize many biologically active compounds with antiviral, antibacterial, antifungal, and even antitumor properties [241] [242] . Perhaps the need for the synthesis of such unusual compounds lies in the absence of reliable protection for the sponges. Sea sponges are the richest source of pharmacologically active compounds among all marine organisms. In 2010, more than 5300 different substances were isolated from the sponges or their symbionts, and more than 200 items are added to this list annually [243] .
An example of compounds synthesized by sponges is the oxidized derivatives of fatty acids , , which have anticancer, antibacterial, and antifungal activity. group compound isolated from sponges, was toxic to mouse lymphoma cells [244] [245] . The sponge synthesizes , a synthetic analogue of which is and has anti - cancer properties [246] . It has been shown that some alkaloids isolated from sponges help protect the kidneys from the cytotoxic effect of the anticancer drug cisplatin [247] . Sponge alkaloids of the genus ( B, C, D) kill the resting forms of the causative agent of tuberculosis Mycobacterium tuberculosis [248] . Nonamphamide B, a cyclic Neamphius sponge, has the same activity [249] . At the same time, , synthesized by the sponge , suppresses HIV [250] . The sponge synthesizes halistanol trisulfate, a broad-spectrum antibiotic that destroys the cytoplasmic membrane [251] . Sponge forms substances with antibacterial and antifungal effects [252] . Spongotimidin and sponguridin isolated from became the prototype for the synthesis of antiviral agents vidarabine , cytarabine and zidovudine [243] . More than 100 substances isolated from sponges and having an antitumor effect, as well as their synthetic analogues and derivatives, were studied in vitro on various tumor cell lines [243] . More than 60 secondary sponge metabolites have antimalarial activity : for example, Axisonitrile-1 from the sponge became the ancestor of isonitrile-containing derivatives and their analogues [243] .
Microorganisms living inside the sponges can serve as natural reservoirs of antibiotic resistance genes, as in the case of the sponge [253] .
Currently, studies are underway on the prospects of using the chitin skeleton of the marine sponges of the Aplysinidae family in the creation of bioengineered structures of adipose , cartilage and bone tissue based on mesenchymal stromal cells [254] [255] .
Powdered dried freshwater sponges of a bug of a bug - a traditional folk remedy for bruises , bruises , rheumatism , which were used to rub the skin, or as an ointment mixed with fat or water. Badyagi medicine was also used as an internal treatment for scrofula [256] [257] [258] . Currently, the drug of the same name is manufactured industrially with local irritating, anti-inflammatory and analgesic effects. Dried budyag powder, despite the harm caused by prolonged use, has also been traditionally used as a cosmetic product and as a cheap substitute for blush [259] . Potters from South America added this powder to pottery clay to increase the strength of its products [260] .
Since 1907, sponges have been used by researchers to study intercellular adhesion, as well as cell and intercellular matrix adhesion [261] . Sponges can serve as convenient objects for researchers, as a model for studying more complex animals [262] . Due to its porous structure, similar to the structure of bone tissue , the presence of spongin - a polymer very similar to animal collagen , widely used in tissue regeneration, and the mineral component, sponges are well suited for use in experiments on creating bone tissue [263] .
Sponges in Culture
The use of sea sponges firmly entered the culture and life of the peoples of the Mediterranean long before the beginning of our era - first of all, as a means of personal hygiene : when bathing and applying olive oil and perfume to the body (in this quality, its skeleton freed from the rest of the body of the sponge, which consists of a network of horny fibers and resembles elastic felt [264] ). The ancient Egyptians used a sponge to embalm the deceased, rubbing his body with aromatic oils. Homer mentions sponges in both of his poems: in the Iliad they give the sponge to Hephaestus , who cleans his hands, face and chest with it, and in the Odyssey Odyssey ’s servants use it to clean the tables in his house after the grooms visit Penelope . Roman soldiers put sponges under their helmets , and doctors used them to clean wounds [265] [266] .
In addition, sponges in ancient times were considered one of the tools used in witchcraft . So, in “ Metamorphoses ” of Apuleius contains a story about how a sorceress killed a man who abandoned her and at the same time used a sponge as a means to delay the moment of death (in order to avert suspicions from a true killer) [267] . A sponge wrapped in silk with a thread tied to it was used by the ancient Jews as a contraceptive [268] .
The need to supply sponges to numerous consumers has led to the development of sponge collection as a special craft . They were so strong and athletically built that diving behind the lips was included in the program of the Olympic Games of antiquity [266] [269] .
The ancient tradition was carried through centuries by the inhabitants of the islands of the Aegean . In the XIX and early XX centuries, the Dodecanese Islands (primarily the island of Kalymnos , in the vicinity of which there were large populations of sponges) turned into the largest center for the extraction and trade of sea sponges [270] . The difficult and often dangerous fishing of divers has become an important topic of local folklore (in particular, many songs ) [271] . Initially, Kalymnos divers jumped overboard almost naked, holding a 15-kilogram flat stone in their hands for quick diving; after the diving suit was invented in 1865, they began to use this new equipment, which allowed to increase the depth and duration of dives. By the end of the 20th century, however, the production and sales of sea sponges decreased significantly, as synthetic sponges displace them [266] [272] .
In addition to Greece , sponge mining off the coast of Syria , Egypt , Libya and Tunisia has a long history, where this trade has also become part of the cultural heritage of the peoples of these countries. The collection of sponges belonging to 5 species of the genera Spongia and Hippospongia : S. officinalis , S. mollissima (it is often, however, included in the species S. officinalis [273] ), , , [268] [274] . In 1841, the production of sea sponges began for export in the Caribbean - first in the Bahamas , somewhat later in Cuba and on the coast of Florida ( Key West ); here, the number of harvested species of sea sponges was supplemented by , , , , and some other species [275] [ 276] .
One of the copies of the toilet sponge , called the , was included in the number of famous Christian relics [277] . According to the Gospel of John , one of the Roman soldiers guarding the crucified Christ , in response to his request to drink, dipped a sponge in vinegar and brought it to Jesus' lips to quench his thirst [278] . The Holy Sponge, which became the object of veneration, was stored first in Jerusalem , then in Constantinople ; its fragments are currently located in four Roman cathedrals ( Lateran Basilica , Basilica of Santa Croce in Jerusalemme , Santa Maria Maggiore and Santa Maria in Trastevere ). Another large fragment of the Holy Sponge was acquired along with several other relics in 1239–1241. French King Louis IX at the Latin Emperor Baldwin II for a huge amount and placed in the Paris Chapel Saint-Chapelle erected by him [277] [279] . A small fragment of the Holy Sponge is also located in the ark of Dionysius , stored in the Moscow Kremlin [280] [281] .
The image of the Holy Sponge often appears in works of art . On the fresco “The Last Judgment ” by Michelangelo (1537-1541), located on the altar wall of the Sistine Chapel in the Vatican , you can see the image of an angel holding a pole with the Holy Sponge at the end [282] [283] . A similar figure of an angel with the Holy Sponge is found in the frescoes by Francesco Primaticcio (1541-1544), decorating the vaults of the choirs of the chapel of St. Mary in Chali ( commune in the Oise department in northern France ) [284] . The sculpture of an angel with the Holy Sponge in his hands became the most famous work of the Italian sculptor , a student of Lorenzo Bernini ; this sculptor, sculpted by Giorgetti in 1668 or 1669 on the instructions of his teacher, is one of ten angel sculptures installed on the bridge of the Holy Angel in Rome (the terracotta head of the future sculpture made by Georgetti as a sketch is stored in the Hermitage [285] ) [286] .
The Holy Sponge (along with the Crown of Thorns taken from the head of the crucified Christ) appears in the lower right corner of Peter Paul Rubens's “ Descent from the Cross ” (1612), which is one of the altar images in the Cathedral of Our Lady of Antwerp [287] .
The skeletons of glass sponges , purified from organic matter, are used as jewelry and souvenirs. The Venus Basket ( Euplectella ) sponge, whose skeleton has the shape of a delicate openwork cylinder with such an intricate structure that it seems to be carved by the hand of a skilled craftsman, is especially appreciated in this plan (in the Victorian era, one copy of this sponge was purchased for a very significant amount of 5 guineas at that time - over 500 pounds in translation into modern money) [288] [289] . Seven hierarchical structural levels were revealed in the skeleton of this sponge, the ordered assembly of which allows one to overcome the natural fragility of glass and give the skeleton considerable mechanical strength [290] .
In Japan, it has long been a custom to give newlyweds a wedding copy of the sponge with a pair of symbionts inside - shrimp of the species (male and female, which enter the sponge by larvae and then live in it all their lives: get out through the small pores of the sponge they can no longer) [170] [291] . Such a gift symbolizes indestructible love and marital fidelity in accordance with the statement “To live together, to be buried together” [292] .
In 1999, Nickelodeon , an American television station, released the animated series SpongeBob SquarePants, directed by S. Hillenberg ; in the series, the characters are representatives of marine fauna, and the main character is a sponge Bob named "Square Pants" [293] . In 2004 and 2015, the channel released full - length animated films ( SpongeBob SquarePants and SpongeBob SquarePants: Sponge from Water , respectively), which continued the animated series [294] .
Notes
- ↑ Dogel V.A. Invertebrate Zoology. 7th ed / Ed. prof. Yu. I. Polyansky . - Ed. 7th, rev. and additional .. - M .: Higher school , 1981. - 606 p. - 50,000 copies. - S. 101.
- ↑ Koltun V.M. Type of Sponge (Porifera, or Spongia) // Life of animals. Volume 1. The simplest. Coelenterates. Worms / Ed. Yu. I. Polyansky , Ch. ed. V. E. Sokolov . - 2nd ed. - M .: Education, 1987. - S. 151. - 448 p.
- ↑ 1 2 3 4 5 6 Zenkevich L.A. History of the system of invertebrates // Guide to Zoology. T. I. Invertebrates: protozoa, sponges, intestinal, flatworms, nemerthins, roundworms, rotifers / Ed. L. A. Zenkevich. - M .; L .: Biomedgiz, 1937 .-- xii + 795 p. - S. 1-49.
- ↑ Osorio Abarzúa C. G. Sobre agentes infecciosos, zoofitos, animálculos e infusorios // Revista chilena de infectologia. - 2007. - Vol. 24, no. 2. - P. 171-174. - DOI : 10.4067 / S0716-10182007000200016 . - PMID 17453079 .
- ↑ 1 2 Ellis J. On the Nature and Formation of Sponges: In a Letter from John Ellis, Esquire, FRS to Dr. Solander, FRS (Eng.) // Philosophical Transactions of the Royal Society of London. - 1765. - 1 January ( vol. 55 ). - P. 280-288 . - ISSN 0261-0523 . - DOI : 10.1098 / rstl.1765.0032 .
- ↑ 1 2 3 Hyman L. H. Metazoa of the Cellular Grade of Construction - Phylum Porifera, the Sponges // The Invertebrares. Vol 1. Protozoa through Ctenophora. - New York: McGraw-Hill Book Company, 1940. - xii + 726 p. - P. 284-364.
- ↑ Grant R. E. Observations and Experiments on the Structure and Functions of the Sponge // Edinburgh Philosophical Journal. - 1825. - Vol. 13. - P. 94-107, 333-346.
- ↑ 1 2 Ereskovsky, 2010 , p. lv.
- ↑ Richards R. J. The Tragic Sense of Life: Ernst Haeckel and the Struggle over Evolutionary Thought . - Chicago: University of Chicago Press, 2008 .-- xx + 512 p. - ISBN 978-0-226-71214-7 . - P. 196-197.
- ↑ Johnston G. A History of British Sponges and Lithophytes . - Edinburgh: John Stark, 1842. - xii + 264 p. - P. 62-65.
- ↑ Scamardella JM Not plants or animals: a brief history of the origin of Kingdoms Protozoa, Protista and Protoctista // International Microbiology. - 1999. - Vol. 2, no. 4. - P. 207-216. - PMID 10943416 .
- ↑ James-Clarke H. Conclusive Proofs of the Animality of the Ciliate Sponges, and of their Affinities with the Infusoria Flagellata // The Annals and Magazine of Natural History. - 1867. - Vol. 19, Series 3. - P. 13-18.
- ↑ Koltun, 1968 , p. 182.
- ↑ Koltun, 1968 , p. 182-183.
- ↑ Ereskovsky, 2010 , p. 235-236, 247.
- ↑ 1 2 Koltun, 1968 , p. 183.
- ↑ Crow W. B. A Synopsis of Biology. 2nd edition . - Elsevier, 2013 .-- xv + 1072 p. - ISBN 978-1-4832-2532-6 . - P. 222, 740.
- ↑ Ereskovsky, 2010 , p. lix.
- ↑ 1 2 Wanninger, 2015 , p. 70.
- ↑ 1 2 Ishijima J. , Iwabe N. , Masuda Y. , Watanabe Y. , Matsuda Y. Sponge cytogenetics - mitotic chromosomes of ten species of freshwater sponge // Zoological Science. - 2008 .-- Vol. 25, no. 5. - P. 480-486. - DOI : 10.2108 / zsj.25.480 . - PMID 18558800 .
- ↑ 1 2 Muller, 2003 , p. four.
- ↑ 1 2 3 4 Yereskovsky, Vishnyakov, 2015 , p. 9-10.
- ↑ Wagner D., Kelley C. D. The largest sponge in the world? // Marine Biodiversity. - 2016 .-- ISSN 1867-1616 . - DOI : 10.1007 / s12526-016-0508-z .
- ↑ Austin W. C., Conway K. W., Barrie J. V., Krautter M. Growth and morphology of a reef-forming glass sponge, Aphrocallistes vastus (Hexactinellida), and implications for recovery from widespread trawl damage // Porifera Research: Biodiversity, Innovation and Sustainability. - Rio de Janeiro Museu Nacional, 2007. - P. 139-145.
- ↑ Ruppert, Fox, Barnes, 2008 , p. 158.
- ↑ 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Animal life . Volume 1. The simplest, intestinal, worms. - M .: Education , 1987 .-- 508 p. - S. 128-153.
- ↑ 1 2 Gage G. D., Tyler P. A. Deep-sea Biology: A Natural History of Organisms at the Deep-Sea Floor . - Cambridge: Cambridge University Press , 1991 .-- xvi + 504 p. - ISBN 0-521-33665-1 . - P. 91-93.
- ↑ 1 2 3 4 Ruppert, Fox, Barnes, 2008 , p. 157-182.
- ↑ Muller, 2003 , p. 3.
- ↑ 1 2 Lavrov AI , Kosevich IA Sponge cell reaggregation: Cellular structure and morphogenetic potencies of multicellular aggregates // Journal of Experimental Zoology. Part A. Ecological Genetics and Physiology. - 2016. - Vol. 325, no. 2. - P. 158-177. - DOI : 10.1002 / jez.2006 . - PMID 26863993 .
- ↑ 1 2 3 Voigt O. , Wülfing E. , Wörheide G. Molecular phylogenetic evaluation of classification and scenarios of character evolution in calcareous sponges (Porifera, Class Calcarea) // PLoS ONE . - 2012. - Vol. 7, no. 3.- P. e33417. - DOI : 10.1371 / journal.pone.0033417 . - PMID 22479395 .
- ↑ 1 2 3 Ruppert, Fox, Barnes, 2008 , p. 148.
- ↑ Yereskovsky, Vishnyakov, 2015 , p. 11-13.
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