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Translocation of noisy miners is unlikely to be a solution to their overabundance in remnant habitats . In a Victorian study where birds were banded and relocated , colonies moved into the now unpopulated area but soon returned to their original territories . The translocated birds did not settle in a new territory . ...
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= Principe Amedeo @-@ class ironclad =
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The Principe Amedeo class was a pair of ironclad warships built for the Italian Regia Marina ( Royal Navy ) in the 1870s and 1880s . They were the culmination of a major naval construction program designed to give Italy a powerful fleet of ironclads . The two ships , Principe Amedeo and Palestro , were the last Italia...
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= = Design = =
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In 1862 , the Italian government under Prime Minister Urbano Rattazzi and his naval minister Carlo Pellion di Persano made the decision to build a fleet of ironclad warships . The Italian fleet had already acquired a pair of small , French @-@ built armored frigates of the Formidabile class , and two more vessels of t...
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= = = General characteristics and machinery = = =
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The two ships differed slightly in size . Principe Amedeo was 79 @.@ 73 meters ( 261 @.@ 6 ft ) long between perpendiculars , while Palestro was 78 @.@ 82 m ( 258 @.@ 6 ft ) long . Principe Amedeo had a beam of 17 @.@ 4 m ( 57 ft ) and a draft of 7 @.@ 9 m ( 26 ft ) ; Palestro 's beam measured 17 @.@ 3 m ( 57 ft ) , a...
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The ships ' propulsion system consisted of one single @-@ expansion steam engine that drove a single screw propeller , with steam supplied by six coal @-@ fired , cylindrical fire @-@ tube boilers . The boilers were trunked into a single funnel . The lead ship 's engine produced a top speed of 12 @.@ 2 knots ( 22 @.@ ...
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= = = Armament and armor = = =
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Palestro and Principe Amedeo were both armed with a main battery of six 10 in ( 254 mm ) guns , though they were mounted differently in each ship . Principe Amedeo carried hers in a single armored casemate located amidships , while Palestro 's guns were mounted in three armored casemates . The first was located forwar...
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The two ships were protected by iron belt armor that was 8 @.@ 7 in ( 221 mm ) thick and extended for the entire length of the hull . The casemates were protected with 5 @.@ 5 in ( 140 mm ) of iron plating , and the small conning tower had 2 @.@ 4 in ( 61 mm ) thick iron plates .
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= = Ships = =
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= = Service history = =
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Neither ship had a particularly eventful career . They were completed too late to take part in the final stages of the wars of Italian unification . Instead , they were assigned to the Italian colonial empire , with occasional stints in the main Italian fleet . In 1880 , Palestro took part in a naval demonstration off...
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In the late 1880s , both ships were withdrawn from frontline service and employed as headquarters ships for the defense of Taranto — Principe Amedeo — and La Maddalena — Palestro . Principe Amedeo was stricken from the naval register in 1895 and used as an ammunition depot ship in Taranto until 1910 , when she was sol...
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= Cell nucleus =
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In cell biology , the nucleus ( pl. nuclei ; from Latin nucleus or nuculeus , meaning kernel ) is a membrane @-@ enclosed organelle found in eukaryotic cells . Eukaryotes usually have a single nucleus , but a few cell types , such as mammalian red blood cells , have no nuclei , and a few others have many .
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Cell nuclei contain most of the cell 's genetic material , organized as multiple long linear DNA molecules in complex with a large variety of proteins , such as histones , to form chromosomes . The genes within these chromosomes are the cell 's nuclear genome and are structured in such a way to promote cell function ....
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Because the nuclear membrane is impermeable to large molecules , nuclear pores are required that regulate nuclear transport of molecules across the envelope . The pores cross both nuclear membranes , providing a channel through which larger molecules must be actively transported by carrier proteins while allowing free...
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= = History = =
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The nucleus was the first organelle to be discovered . What is most likely the oldest preserved drawing dates back to the early microscopist Antonie van Leeuwenhoek ( 1632 – 1723 ) . He observed a " lumen " , the nucleus , in the red blood cells of salmon . Unlike mammalian red blood cells , those of other vertebrates...
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The nucleus was also described by Franz Bauer in 1804 and in more detail in 1831 by Scottish botanist Robert Brown in a talk at the Linnean Society of London . Brown was studying orchids under microscope when he observed an opaque area , which he called the " areola " or " nucleus " , in the cells of the flower 's out...
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He did not suggest a potential function . In 1838 , Matthias Schleiden proposed that the nucleus plays a role in generating cells , thus he introduced the name " cytoblast " ( cell builder ) . He believed that he had observed new cells assembling around " cytoblasts " . Franz Meyen was a strong opponent of this view ,...
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Between 1877 and 1878 , Oscar Hertwig published several studies on the fertilization of sea urchin eggs , showing that the nucleus of the sperm enters the oocyte and fuses with its nucleus . This was the first time it was suggested that an individual develops from a ( single ) nucleated cell . This was in contradictio...
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= = Structures = =
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The nucleus is the largest cellular organelle in animal cells . In mammalian cells , the average diameter of the nucleus is approximately 6 micrometres ( µm ) , which occupies about 10 % of the total cell volume . The viscous liquid within it is called nucleoplasm , and is similar in composition to the cytosol found o...
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= = = Nuclear envelope and pores = = =
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The nuclear envelope , otherwise known as nuclear membrane , consists of two cellular membranes , an inner and an outer membrane , arranged parallel to one another and separated by 10 to 50 nanometres ( nm ) . The nuclear envelope completely encloses the nucleus and separates the cell 's genetic material from the surr...
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Nuclear pores , which provide aqueous channels through the envelope , are composed of multiple proteins , collectively referred to as nucleoporins . The pores are about 125 million daltons in molecular weight and consist of around 50 ( in yeast ) to several hundred proteins ( in vertebrates ) . The pores are 100 nm in...
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Most proteins , ribosomal subunits , and some DNAs are transported through the pore complexes in a process mediated by a family of transport factors known as karyopherins . Those karyopherins that mediate movement into the nucleus are also called importins , whereas those that mediate movement out of the nucleus are c...
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= = = Nuclear lamina = = =
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In animal cells , two networks of intermediate filaments provide the nucleus with mechanical support : The nuclear lamina forms an organized meshwork on the internal face of the envelope , while less organized support is provided on the cytosolic face of the envelope . Both systems provide structural support for the n...
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The nuclear lamina is composed mostly of lamin proteins . Like all proteins , lamins are synthesized in the cytoplasm and later transported to the nucleus interior , where they are assembled before being incorporated into the existing network of nuclear lamina . Lamins found on the cytosolic face of the membrane , suc...
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Like the components of other intermediate filaments , the lamin monomer contains an alpha @-@ helical domain used by two monomers to coil around each other , forming a dimer structure called a coiled coil . Two of these dimer structures then join side by side , in an antiparallel arrangement , to form a tetramer calle...
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Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as laminopathies . The most notable laminopathy is the family of diseases known as progeria , which causes the appearance of premature aging in its sufferers . The exact mechanism by which the associated bioc...
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= = = Chromosomes = = =
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The cell nucleus contains the majority of the cell 's genetic material in the form of multiple linear DNA molecules organized into structures called chromosomes . Each human cell contains roughly two meters of DNA . During most of the cell cycle these are organized in a DNA @-@ protein complex known as chromatin , and...
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There are two types of chromatin . Euchromatin is the less compact DNA form , and contains genes that are frequently expressed by the cell . The other type , heterochromatin , is the more compact form , and contains DNA that is infrequently transcribed . This structure is further categorized into facultative heterochr...
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Antibodies to certain types of chromatin organization , in particular , nucleosomes , have been associated with a number of autoimmune diseases , such as systemic lupus erythematosus . These are known as anti @-@ nuclear antibodies ( ANA ) and have also been observed in concert with multiple sclerosis as part of gener...
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= = = Nucleolus = = =
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The nucleolus is a discrete densely stained structure found in the nucleus . It is not surrounded by a membrane , and is sometimes called a suborganelle . It forms around tandem repeats of rDNA , DNA coding for ribosomal RNA ( rRNA ) . These regions are called nucleolar organizer regions ( NOR ) . The main roles of th...
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In the first step of ribosome assembly , a protein called RNA polymerase I transcribes rDNA , which forms a large pre @-@ rRNA precursor . This is cleaved into the subunits 5.8S , 18S , and 28S rRNA . The transcription , post @-@ transcriptional processing , and assembly of rRNA occurs in the nucleolus , aided by smal...
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When observed under the electron microscope , the nucleolus can be seen to consist of three distinguishable regions : the innermost fibrillar centers ( FCs ) , surrounded by the dense fibrillar component ( DFC ) , which in turn is bordered by the granular component ( GC ) . Transcription of the rDNA occurs either in t...
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= = = Other subnuclear bodies = = =
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Besides the nucleolus , the nucleus contains a number of other non @-@ membrane @-@ delineated bodies . These include Cajal bodies , Gemini of coiled bodies , polymorphic interphase karyosomal association ( PIKA ) , promyelocytic leukaemia ( PML ) bodies , paraspeckles , and splicing speckles . Although little is know...
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Other subnuclear structures appear as part of abnormal disease processes . For example , the presence of small intranuclear rods has been reported in some cases of nemaline myopathy . This condition typically results from mutations in actin , and the rods themselves consist of mutant actin as well as other cytoskeleta...
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= = = = Cajal bodies and gems = = = =
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A nucleus typically contains between 1 and 10 compact structures called Cajal bodies or coiled bodies ( CB ) , whose diameter measures between 0 @.@ 2 µm and 2 @.@ 0 µm depending on the cell type and species . When seen under an electron microscope , they resemble balls of tangled thread and are dense foci of distribu...
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Similar to Cajal bodies are Gemini of Cajal bodies , or gems , whose name is derived from the Gemini constellation in reference to their close " twin " relationship with CBs . Gems are similar in size and shape to CBs , and in fact are virtually indistinguishable under the microscope . Unlike CBs , gems do not contain...
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= = = = RAFA and PTF domains = = = =
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RAFA domains , or polymorphic interphase karyosomal associations , were first described in microscopy studies in 1991 . Their function remains unclear , though they were not thought to be associated with active DNA replication , transcription , or RNA processing . They have been found to often associate with discrete ...
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= = = = PML bodies = = = =
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Promyelocytic leukaemia bodies ( PML bodies ) are spherical bodies found scattered throughout the nucleoplasm , measuring around 0 @.@ 1 – 1 @.@ 0 µm . They are known by a number of other names , including nuclear domain 10 ( ND10 ) , Kremer bodies , and PML oncogenic domains . PML bodies are named after one of their ...
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= = = = Splicing speckles = = = =
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Speckles are subnuclear structures that are enriched in pre @-@ messenger RNA splicing factors and are located in the interchromatin regions of the nucleoplasm of mammalian cells . At the fluorescence @-@ microscope level they appear as irregular , punctate structures , which vary in size and shape , and when examined...
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= = = = Paraspeckles = = = =
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Discovered by Fox et al. in 2002 , paraspeckles are irregularly shaped compartments in the nucleus ' interchromatin space . First documented in HeLa cells , where there are generally 10 – 30 per nucleus , paraspeckles are now known to also exist in all human primary cells , transformed cell lines , and tissue sections...
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Paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity . They are transcription dependent and in the absence of RNA Pol II transcription , the paraspeckle disappears and all of its associated protein components ( PSP1 , p54nrb , PSP2 , CFI ( m ) 68 , and PSF ) form a...
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= = = = Perichromatin fibrils = = = =
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Perichromatin fibrils are visible only under electron microscope . They are located next to the transcriptionally active chromatin and are hypothesized to be the sites of active pre @-@ mRNA processing .
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= = Function = =
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The nucleus provides a site for genetic transcription that is segregated from the location of translation in the cytoplasm , allowing levels of gene regulation that are not available to prokaryotes . The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell ...
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The nucleus is an organelle found in eukaryotic cells . Inside its fully enclosed nuclear membrane , it contains the majority of the cell 's genetic material . This material is organized as DNA molecules , along with a variety of proteins , to form chromosomes .
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= = = Cell compartmentalization = = =
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The nuclear envelope allows the nucleus to control its contents , and separate them from the rest of the cytoplasm where necessary . This is important for controlling processes on either side of the nuclear membrane . In most cases where a cytoplasmic process needs to be restricted , a key participant is removed to th...
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In order to control which genes are being transcribed , the cell separates some transcription factor proteins responsible for regulating gene expression from physical access to the DNA until they are activated by other signaling pathways . This prevents even low levels of inappropriate gene expression . For example , ...
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The compartmentalization allows the cell to prevent translation of unspliced mRNA . Eukaryotic mRNA contains introns that must be removed before being translated to produce functional proteins . The splicing is done inside the nucleus before the mRNA can be accessed by ribosomes for translation . Without the nucleus ,...
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= = = Gene expression = = =
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Gene expression first involves transcription , in which DNA is used as a template to produce RNA . In the case of genes encoding proteins , that RNA produced from this process is messenger RNA ( mRNA ) , which then needs to be translated by ribosomes to form a protein . As ribosomes are located outside the nucleus , m...
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Since the nucleus is the site of transcription , it also contains a variety of proteins that either directly mediate transcription or are involved in regulating the process . These proteins include helicases , which unwind the double @-@ stranded DNA molecule to facilitate access to it , RNA polymerases , which synthe...
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= = = Processing of pre @-@ mRNA = = =
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Newly synthesized mRNA molecules are known as primary transcripts or pre @-@ mRNA . They must undergo post @-@ transcriptional modification in the nucleus before being exported to the cytoplasm ; mRNA that appears in the cytoplasm without these modifications is degraded rather than used for protein translation . The t...
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RNA splicing , carried out by a complex called the spliceosome , is the process by which introns , or regions of DNA that do not code for protein , are removed from the pre @-@ mRNA and the remaining exons connected to re @-@ form a single continuous molecule . This process normally occurs after 5 ' capping and 3 ' po...
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= = Dynamics and regulation = =
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= = = Nuclear transport = = =
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The entry and exit of large molecules from the nucleus is tightly controlled by the nuclear pore complexes . Although small molecules can enter the nucleus without regulation , macromolecules such as RNA and proteins require association karyopherins called importins to enter the nucleus and exportins to exit . " Cargo...
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Nuclear import depends on the importin binding its cargo in the cytoplasm and carrying it through the nuclear pore into the nucleus . Inside the nucleus , RanGTP acts to separate the cargo from the importin , allowing the importin to exit the nucleus and be reused . Nuclear export is similar , as the exportin binds th...
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Specialized export proteins exist for translocation of mature mRNA and tRNA to the cytoplasm after post @-@ transcriptional modification is complete . This quality @-@ control mechanism is important due to these molecules ' central role in protein translation . Mis @-@ expression of a protein due to incomplete excisio...
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= = = Assembly and disassembly = = =
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During its lifetime , a nucleus may be broken down or destroyed , either in the process of cell division or as a consequence of apoptosis ( the process of programmed cell death ) . During these events , the structural components of the nucleus — the envelope and lamina — can be systematically degraded . In most cells ...
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At a certain point during the cell cycle in open mitosis , the cell divides to form two cells . In order for this process to be possible , each of the new daughter cells must have a full set of genes , a process requiring replication of the chromosomes as well as segregation of the separate sets . This occurs by the r...
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However , in dinoflagellates , the nuclear envelope remains intact , the centrosomes are located in the cytoplasm , and the microtubules come in contact with chromosomes , whose centromeric regions are incorporated into the nuclear envelope ( the so @-@ called closed mitosis with extranuclear spindle ) . In many other...
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Apoptosis is a controlled process in which the cell 's structural components are destroyed , resulting in death of the cell . Changes associated with apoptosis directly affect the nucleus and its contents , for example , in the condensation of chromatin and the disintegration of the nuclear envelope and lamina . The d...
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The nuclear envelope acts as a barrier that prevents both DNA and RNA viruses from entering the nucleus . Some viruses require access to proteins inside the nucleus in order to replicate and / or assemble . DNA viruses , such as herpesvirus replicate and assemble in the cell nucleus , and exit by budding through the i...
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= = = Disease @-@ related dynamics = = =
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Initially , it has been suspected that immunoglobulins in general and autoantibodies in particular do not enter the nucleus . Now there is a body of evidence that under pathological conditions ( e.g. lupus erythematosus ) IgG can enter the nucleus .
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= = Nuclei per cell = =
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Most eukaryotic cell types usually have a single nucleus , but some have no nuclei , while others have several . This can result from normal development , as in the maturation of mammalian red blood cells , or from faulty cell division .
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= = = Anucleated cells = = =
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An anucleated cell contains no nucleus and is , therefore , incapable of dividing to produce daughter cells . The best @-@ known anucleated cell is the mammalian red blood cell , or erythrocyte , which also lacks other organelles such as mitochondria , and serves primarily as a transport vessel to ferry oxygen from th...
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In flowering plants , this condition occurs in sieve tube elements .
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= = = Multinucleated cells = = =
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Multinucleated cells contain multiple nuclei . Most acantharean species of protozoa and some fungi in mycorrhizae have naturally multinucleated cells . Other examples include the intestinal parasites in the genus Giardia , which have two nuclei per cell . In humans , skeletal muscle cells , called myocytes and syncyti...
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A number of dinoflagelates are known to have two nuclei . Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA : one from the dinoflagelate and the other from a symbiotic diatom . Curiously the mitochondrion and the plastid of the diatom remain functional .
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= = Evolution = =
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As the major defining characteristic of the eukaryotic cell , the nucleus ' evolutionary origin has been the subject of much speculation . Four major hypotheses have been proposed to explain the existence of the nucleus , although none have yet earned widespread support .
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The first model known as the " syntrophic model " proposes that a symbiotic relationship between the archaea and bacteria created the nucleus @-@ containing eukaryotic cell . ( Organisms of the Archaea and Bacteria domain have no cell nucleus . ) It is hypothesized that the symbiosis originated when ancient archaea , ...
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A second model proposes that proto @-@ eukaryotic cells evolved from bacteria without an endosymbiotic stage . This model is based on the existence of modern planctomycetes bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures . A similar proposal states that a ...
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The most controversial model , known as viral eukaryogenesis , posits that the membrane @-@ bound nucleus , along with other eukaryotic features , originated from the infection of a prokaryote by a virus . The suggestion is based on similarities between eukaryotes and viruses such as linear DNA strands , mRNA capping ...
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A more recent proposal , the exomembrane hypothesis , suggests that the nucleus instead originated from a single ancestral cell that evolved a second exterior cell membrane ; the interior membrane enclosing the original cell then became the nuclear membrane and evolved increasingly elaborate pore structures for passag...
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