Property Summary

NCBI Gene PubMed Count 126
PubMed Score 48.34
PubTator Score 660.23

Knowledge Summary


No data available


  Differential Expression (26)

Disease log2 FC p
pancreatic cancer 1.900 1.7e-03
astrocytic glioma -2.100 4.9e-02
oligodendroglioma -1.900 1.7e-02
chronic lymphosyte leukemia -1.300 9.5e-09
glioblastoma 2.100 1.0e-02
medulloblastoma, large-cell -1.700 2.5e-04
Duchenne muscular dystrophy 1.869 6.8e-11
autosomal dominant Emery-Dreifuss muscul... 1.630 2.2e-03
Becker muscular dystrophy 1.006 1.8e-02
juvenile dermatomyositis 1.746 2.6e-11
limb girdle muscular dystrophy 2A 1.016 3.3e-04
acute quadriplegic myopathy 1.321 3.4e-05
primary pancreatic ductal adenocarcinoma 1.722 1.5e-03
Common variable immunodeficiency -1.526 2.4e-02
colon cancer -1.200 1.2e-02
lung cancer 1.200 2.2e-03
group 4 medulloblastoma -1.600 9.5e-04
posterior fossa group A ependymoma 1.200 3.8e-05
pancreatic carcinoma 1.200 1.6e-04
subependymal giant cell astrocytoma 2.214 3.2e-02
lung adenocarcinoma 1.172 4.0e-06
COPD -1.100 3.1e-04
gastric carcinoma 2.000 1.6e-02
ulcerative colitis 1.200 5.2e-08
ovarian cancer 3.200 3.3e-07
dermatomyositis 1.600 1.2e-02

 CSPA Cell Line (3)

Protein-protein Interaction (4)

Gene RIF (48)

26879394 Several dominantly inherited congenital macrothrombocytopenia-associated mutations that are located within the actinin-1 actin-binding domain cause increased binding of actinin-1 to actin filaments and enhance filament bundling in vitro.
26854759 The authors show that the microtubule binding protein EB1 (end-binding protein 1), a key regulator of microtubule dynamics, can bind directly to filamentous actin (F-actin) F-actin.
26609810 Drp1 is in dynamic equilibrium on mitochondria in a fission-independent manner, and that fission factors such as actin filaments target productive oligomerization to fission sites.
26096820 Host cortactin, PKD1 and actin are recruited by Trypanosoma cruzi extracellular amastigotes based on experiments in fixed and live cells by time lapse confocal microscopy.
26071203 Data indicate that alpha-actinin (110 kDa) autoantibody was characterized as a major target in lupus nephritis (LN) patients positive for Anti-membrane autoantibodies (MbA).
25431851 ACTN1 determines the motility of keratinocytes by regulating the organization of the actin cytoskeleton, focal adhesion, and hemidesmosome proteins complexes, thereby modulating cell speed, lamellipodial dynamics, and directed migration
25429109 ArgBP2 interaction with alpha-actinin and actin stress fibers inhibits cell migration
25361813 ACTN1-related thrombocytopenia is characterized by mild course with platelet macrocytosis and low risk for bleeding.
24069336 A missense mutation in the alpha-actinin 1 gene (ACTN1) is the cause of autosomal dominant macrothrombocytopenia in a large French family.
23664615 We conclude that alpha-actinin stabilizes Ca(V)1.2 at the plasma membrane and that its displacement by calcium-calmodulin triggers calcium-induced endocytosis of Ca(V)1.2, thus providing an important negative feedback mechanism for calcium influx.
23557398 Actinin-alpha1 readily forms heterodimers composed of monomers that may have different properties and interacting proteins, altering our view of non-muscle actinin, thus altering our view of non-muscle actinin function.
23454549 The tyrosine phosphorylation of alpha-actinin1 at Y12 and alpha-actinin4 at Y265 is critical for dorsal stress fiber establishment, transverse arc maintenance and focal adhesion maturation.
23434115 Transduction of mouse fetal liver-derived megakaryocytes with disease-associated ACTN1 variants caused a disorganized actin-based cytoskeleton in megakaryocytes.
23139387 Alpha actinin 1, a cytoskeleton protein implicated in inflammatory/degenerative autoimmune diseases, might be regarded as a novel multiple sclerosis autoantigen.
23132927 Loss of dorsal stress fibers either by depleting alpha-actinin-1 or Rac1 results in a beta-actin accumulation at the leading edge in migrating and spreading cells.
22689882 The alpha-actinin/EWI motif-containing protein 2 (EWI-2) complex is involved in regulation of the actin cytoskeleton at T cell immune and virological synapses, providing a link between membrane microdomains and structures involved in T cell recognition.
22659138 The C-terminal polybasic region of CYTH2 participates in interacting directly with the EFh2 domain of ACTN1.
21474972 Actinin-1 and cortactin showed matrix-contact-side localization in adenocarcinoma cells.
21408150 The results suggest that the interaction between HAMLET and alpha-actinins promotes tumor cell detachment.
20634891 Observational study of gene-disease association. (HuGE Navigator)
20156433 Alpha-actinin 1 and 4 are differentially regulated during the development and progression of astrocytomas because each of these isoforms uniquely contributes to distinct malignant properties of astrocytoma cells.
20037648 Both alpha-actinin-1 and alpha-actinin-4 make critical and distinct contributions to cytoskeletal organization, rigidity-sensing, and motility of glioma cells.
19750481 cathepsin X overexpression reduced LFA-1 clustering and induced an intermediate affinity LFA-1 conformation that is known to associate with alpha-actinin-1.
19715605 Mactinin is a novel inducer of Hsp90 activity on monocytes and may serve to perpetuate and augment monocytic activation.
19094982 Data describe alterations of myocardial intercellular and cell-matrix contacts in hypertrophic tissue, and show intracellular translocation of beta-catenin, alpha-actinin and chondroitin sulfate proteoglycan 6 in both an animal model and in LVH patients.
18978946 ICAM-2 mediates suppression of metastatic phenotype and the interaction of ICAM-2/alpha-actinin/actin represents the first complete membrane-linker protein-actin linkage to impact tumor cell motility in vitro and metastatic potential in an in vivo model.
18560563 interaction of GNE with alpha-actinin 1 might point to its involvement in alpha-actinin mediated processes
18519573 two sm-titin Zq domains interact with each other and with the two R2-R3 sites in the alpha-actinin central rod
18408146 alpha-actinin-1 may play a role in human glomerular disease
18353764 A subset of the tumor-specific splicing alterations (ACTN1, CALD1, and VCL) was found in all three organs and may represent general cancer-related splicing events.
17965186 study examines the mechanism by which phosphoinositide binding regulates alpha-actinin function
17944866 Taken together, alpha-actinin not only attaches TRPP3 to the cytoskeleton but also up-regulates TRPP3 channel function.
17898132 Alpha-actinin-1 phosphorylation at Y12 plays a crucial role in pressure-activated cell adhesion and mechanotransduction by facilitating Src recruitment to beta(1)-integrin, and consequently the association of focal adhesion kinase with Src.
17525629 ACT1 is de-expressed in endometriosis and endometrioid carcinoma compared with normal uterine epithelium.
17311919 there is an alpha-actinin-1-dependent mGlu(5b) receptor association with the actin cytoskeleton modulating receptor cell surface expression and functioning
17115122 Molecular dynamics method was applied to investigate the mechanical behaviour of the human skeletal muscle alpha-actinin.
16951376 Interaction of alpha 1 actinin with ICAM-1 is required for leukocyte extravasation.
16820411 Results show that the interaction between ICAM-5 and alpha-actinin is mediated through binding of positively charged amino acids near the transmembrane domain of ICAM-5, and this interaction may play an important role in neuronal differentiation.
16807302 Alpha-actinin plays a role in regulating cell survival through stabilization of focal adhesions and regulation of TNF-alpha-induced survival signaling.
16430917 Results show that the interaction between two species with weak affinity in solution, such as vinculin and alpha-actinin, can be visualized in a membrane-like environment.
16195373 S1P-induced recruitment of S1P1 to CEM fractions promotes PI3 kinase-mediated Tiam1/Rac1 activation required for alpha-actinin-1/4-regulated cortical actin rearrangement and EC barrier enhancement
16190873 GluR4 may regulate its synaptic targeting through phosphorylation-dependent interactions with alpha-Actinin-1 and IQGAP1
15710624 the dynamics of alpha-actinin are important for PI 3-kinase-induced reorganization of the actin cytoskeleton
12837758 an alpha-actinin-dependent association between the actin cytoskeleton and A2AR trafficking
12493766 the focal adhesion component alpha-actinin interacts with syndecan-4 in a beta-integrin-independent manner
10029173 Identification of a 31 kD amino-terminal fragment of alpha-actinin protein, named mactinin, that promotes monocyte/macrophage maturation in vitro.
8997639 Alpha-actinin is cleaved in vitro by HIV-1 protease at amino acid residues 166-167 and 471-472
1991513 Alpha-actinin is cleaved in vitro by HIV-1 protease at amino acid residues 166-167 and 471-472

AA Sequence


Text Mined References (133)

PMID Year Title
26879394 2016 Congenital macrothrombocytopenia-linked mutations in the actin-binding domain of ?-actinin-1 enhance F-actin association.
26854759 2016 Interactions between the Microtubule Binding Protein EB1 and F-Actin.
26609810 2015 Actin filaments target the oligomeric maturation of the dynamin GTPase Drp1 to mitochondrial fission sites.
26096820 2015 Trypanosoma cruzi extracellular amastigotes trigger the protein kinase D1-cortactin-actin pathway during cell invasion.
26071203 2015 Anti-alpha-actinin antibodies are part of the anti-cell membrane antibody spectrum that characterize patients with lupus nephritis.
25944712 2015 N-terminome analysis of the human mitochondrial proteome.
25431851 2015 Alpha actinin-1 regulates cell-matrix adhesion organization in keratinocytes: consequences for skin cell motility.
25429109 2015 Arg kinase-binding protein 2 (ArgBP2) interaction with ?-actinin and actin stress fibers inhibits cell migration.
25416956 2014 A proteome-scale map of the human interactome network.
25361813 2015 ACTN1-related thrombocytopenia: identification of novel families for phenotypic characterization.
25241761 2014 Using an in situ proximity ligation assay to systematically profile endogenous protein-protein interactions in a pathway network.
24275569 2014 An enzyme assisted RP-RPLC approach for in-depth analysis of human liver phosphoproteome.
24069336 2013 A missense mutation in the alpha-actinin 1 gene (ACTN1) is the cause of autosomal dominant macrothrombocytopenia in a large French family.
24024966 2013 Genome-wide association study of chronic periodontitis in a general German population.
23969696 2013 Multiethnic meta-analysis of genome-wide association studies in >100 000 subjects identifies 23 fibrinogen-associated Loci but no strong evidence of a causal association between circulating fibrinogen and cardiovascular disease.
23664615 2013 Competition between ?-actinin and Ca²?-calmodulin controls surface retention of the L-type Ca²? channel Ca(V)1.2.
23557398 2013 An analysis of splicing, actin-binding properties, heterodimerization and molecular interactions of the non-muscle ?-actinins.
23535732 2013 Identification of 23 new prostate cancer susceptibility loci using the iCOGS custom genotyping array.
23533145 2013 In-depth proteomic analyses of exosomes isolated from expressed prostatic secretions in urine.
23454549 2013 ?-actinin1 and 4 tyrosine phosphorylation is critical for stress fiber establishment, maintenance and focal adhesion maturation.
23434115 2013 ACTN1 mutations cause congenital macrothrombocytopenia.
23139387 2013 Alpha actinin is specifically recognized by Multiple Sclerosis autoantibodies isolated using an N-glucosylated peptide epitope.
23132927 2013 Assembly of non-contractile dorsal stress fibers requires ?-actinin-1 and Rac1 in migrating and spreading cells.
22689882 2012 EWI-2 association with ?-actinin regulates T cell immune synapses and HIV viral infection.
22659138 2012 Arf6 guanine-nucleotide exchange factor, cytohesin-2, interacts with actinin-1 to regulate neurite extension.
22351778 2012 Familial focal segmental glomerulosclerosis (FSGS)-linked ?-actinin 4 (ACTN4) protein mutants lose ability to activate transcription by nuclear hormone receptors.
21784188 2011 ?-Actinin-3 deficiency is associated with reduced bone mass in human and mouse.
21474972 2011 Localization of the invadopodia-related proteins actinin-1 and cortactin to matrix-contact-side cytoplasm of cancer cells in surgically resected lung adenocarcinomas.
21423176 2011 Analysis of the myosin-II-responsive focal adhesion proteome reveals a role for ?-Pix in negative regulation of focal adhesion maturation.
21408150 2011 HAMLET binding to ?-actinin facilitates tumor cell detachment.
21406692 2011 System-wide temporal characterization of the proteome and phosphoproteome of human embryonic stem cell differentiation.
21362503 2011 Protein profile of exosomes from trabecular meshwork cells.
21269460 2011 Initial characterization of the human central proteome.
21266579 2011 Raftlin is involved in the nucleocapture complex to induce poly(I:C)-mediated TLR3 activation.
20634891 2010 Maternal genes and facial clefts in offspring: a comprehensive search for genetic associations in two population-based cleft studies from Scandinavia.
20156433 2010 Alpha-actinin 1 and alpha-actinin 4: contrasting roles in the survival, motility, and RhoA signaling of astrocytoma cells.
20037648 2009 Isoform-specific contributions of alpha-actinin to glioma cell mechanobiology.
19943616 2010 Cell membrane expression of cardiac sodium channel Na(v)1.5 is modulated by alpha-actinin-2 interaction.
19750481 2009 Cathepsin X cleaves the beta2 cytoplasmic tail of LFA-1 inducing the intermediate affinity form of LFA-1 and alpha-actinin-1 binding.
19715605 2009 Mactinin, a fragment of cytoskeletal alpha-actinin, is a novel inducer of heat shock protein (Hsp)-90 mediated monocyte activation.
19608861 2009 Lysine acetylation targets protein complexes and co-regulates major cellular functions.
19247474 2009 Genome-wide and candidate gene association study of cigarette smoking behaviors.
19199708 2009 Proteomic analysis of human parotid gland exosomes by multidimensional protein identification technology (MudPIT).
19094982 2009 Expression and tissue localization of beta-catenin, alpha-actinin and chondroitin sulfate proteoglycan 6 is modulated during rat and human left ventricular hypertrophy.
19056867 2009 Large-scale proteomics and phosphoproteomics of urinary exosomes.
18978946 2008 ICAM-2 expression mediates a membrane-actin link, confers a nonmetastatic phenotype and reflects favorable tumor stage or histology in neuroblastoma.
18560563 2008 UDP-N-acetylglucosamine 2-epimerase/N-acetylmannosamine kinase (GNE) binds to alpha-actinin 1: novel pathways in skeletal muscle?
18519573 2008 Smooth muscle titin Zq domain interaction with the smooth muscle alpha-actinin central rod.
18408146 2008 Expression of alpha-actinin-1 in human glomerular mesangial cells in vivo and in vitro.
18353764 2008 Alternative splicing in colon, bladder, and prostate cancer identified by exon array analysis.
18029348 2008 Toward a confocal subcellular atlas of the human proteome.
17965186 2007 Phosphoinositide binding regulates alpha-actinin CH2 domain structure: analysis by hydrogen/deuterium exchange mass spectrometry.
17944866 2007 Direct binding of alpha-actinin enhances TRPP3 channel activity.
17898132 2007 Alpha-actinin-1 phosphorylation modulates pressure-induced colon cancer cell adhesion through regulation of focal adhesion kinase-Src interaction.
17525629 2007 The cytoskeletal proteins alpha-actinin, Ezrin, and talin are De-expressed in endometriosis and endometrioid carcinoma compared with normal uterine epithelium.
17311919 2007 Actin-binding protein alpha-actinin-1 interacts with the metabotropic glutamate receptor type 5b and modulates the cell surface expression and function of the receptor.
17115122 2007 Mechanical response and conformational changes of alpha-actinin domains during unfolding: a molecular dynamics study.
17081983 2006 Global, in vivo, and site-specific phosphorylation dynamics in signaling networks.
16951376 2006 Evidence of a functional role for interaction between ICAM-1 and nonmuscle alpha-actinins in leukocyte diapedesis.
16820411 2006 alpha-Actinin-dependent cytoskeletal anchorage is important for ICAM-5-mediated neuritic outgrowth.
16807302 2006 Disruption of alpha-actinin-integrin interactions at focal adhesions renders osteoblasts susceptible to apoptosis.
16713569 2006 A protein-protein interaction network for human inherited ataxias and disorders of Purkinje cell degeneration.
16698282 2006 Crystal structure of the actin-binding domain of alpha-actinin 1: evaluating two competing actin-binding models.
16502470 2006 Human colostrum: identification of minor proteins in the aqueous phase by proteomics.
16464232 2006 Postsynaptic recruitment of Dendrin depends on both dendritic mRNA transport and synaptic anchoring.
16430917 2006 Structure of the alpha-actinin-vinculin head domain complex determined by cryo-electron microscopy.
16396496 2006 Insulin-dependent interactions of proteins with GLUT4 revealed through stable isotope labeling by amino acids in cell culture (SILAC).
16291744 2006 Phosphorylated alpha-actinin and protein-tyrosine phosphatase 1B coregulate the disassembly of the focal adhesion kinase x Src complex and promote cell migration.
16195373 2005 Regulation of sphingosine 1-phosphate-induced endothelial cytoskeletal rearrangement and barrier enhancement by S1P1 receptor, PI3 kinase, Tiam1/Rac1, and alpha-actinin.
16190873 2005 Phosphorylation-dependent interactions of alpha-Actinin-1/IQGAP1 with the AMPA receptor subunit GluR4.
16189514 2005 Towards a proteome-scale map of the human protein-protein interaction network.
16097034 2005 Global phosphoproteome analysis on human HepG2 hepatocytes using reversed-phase diagonal LC.
15988023 2005 Structural dynamics of alpha-actinin-vinculin interactions.
15710624 2005 Phosphoinositide binding regulates alpha-actinin dynamics: mechanism for modulating cytoskeletal remodeling.
15489334 2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC).
15159419 2004 Affixin interacts with alpha-actinin and mediates integrin signaling for reorganization of F-actin induced by initial cell-substrate interaction.
15070900 2004 The protein-tyrosine phosphatase SHP-1 regulates the phosphorylation of alpha-actinin.
14702039 2004 Complete sequencing and characterization of 21,243 full-length human cDNAs.
14630798 2004 Characterization of the proteins released from activated platelets leads to localization of novel platelet proteins in human atherosclerotic lesions.
14623081 2003 Direct interaction between alpha-actinin and hepatitis C virus NS5B.
12837758 2003 The adenosine A2A receptor interacts with the actin-binding protein alpha-actinin.
12695497 2003 The conformational state of Tes regulates its zyxin-dependent recruitment to focal adhesions.
12665801 2003 Exploring proteomes and analyzing protein processing by mass spectrometric identification of sorted N-terminal peptides.
12642359 2003 Mutations in the human muscle LIM protein gene in families with hypertrophic cardiomyopathy.
12615977 2003 The lipoma preferred partner LPP interacts with alpha-actinin.
12508121 2003 The DNA sequence and analysis of human chromosome 14.
12507293 2003 Immunohistochemical localization of human fructose-1,6-bisphosphatase in subcellular structures of myocytes.
12499399 2003 Myotilin, the limb-girdle muscular dystrophy 1A (LGMD1A) protein, cross-links actin filaments and controls sarcomere assembly.
12493766 2003 Syndecan-4 associates with alpha-actinin.
12477932 2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences.
12446711 2003 ADIP, a novel Afadin- and alpha-actinin-binding protein localized at cell-cell adherens junctions.
12223541 2002 The cyclin-dependent kinase 5 activators p35 and p39 interact with the alpha-subunit of Ca2+/calmodulin-dependent protein kinase II and alpha-actinin-1 in a calcium-dependent manner.
12211109 2002 Amorphin is phosphorylase; phosphorylase is an alpha-actinin-binding protein.
11973348 2002 Clik1: a novel kinase targeted to actin stress fibers by the CLP-36 PDZ-LIM protein.
11972044 2002 Functional consequences of caspase activation in cardiac myocytes.
11854440 2002 Intracellular domains of NR2 alter calcium-dependent inactivation of N-methyl-D-aspartate receptors.
11846417 2002 Titin mutations as the molecular basis for dilated cardiomyopathy.
11842093 2002 Calsarcin-3, a novel skeletal muscle-specific member of the calsarcin family, interacts with multiple Z-disc proteins.
11802708 2002 Pathological shear stress stimulates the tyrosine phosphorylation of alpha-actinin associated with the glycoprotein Ib-IX complex.
11739652 2001 Interactions of a hemidesmosome component and actinin family members.
11724819 2001 Raver1, a dual compartment protein, is a ligand for PTB/hnRNPI and microfilament attachment proteins.
11573089 2001 Ca2+-independent binding of an EF-hand domain to a novel motif in the alpha-actinin-titin complex.
11470434 2001 Crystal structure of the alpha-actinin rod reveals an extensive torsional twist.
11423549 2001 Analysis of the alpha-actinin/zyxin interaction.
11369769 2001 The cytoskeletal/non-muscle isoform of alpha-actinin is phosphorylated on its actin-binding domain by the focal adhesion kinase.
11305911 2001 A structural characterization of the interactions between titin Z-repeats and the alpha-actinin C-terminal domain.
11223950 2001 Dynamics of alpha-actinin in focal adhesions and stress fibers visualized with alpha-actinin-green fluorescent protein.
11114196 2000 Calsarcins, a novel family of sarcomeric calcineurin-binding proteins.
11113628 2000 Modification of the composition of polycystin-1 multiprotein complexes by calcium and tyrosine phosphorylation.
11110697 2000 Human CLP36, a PDZ-domain and LIM-domain protein, binds to alpha-actinin-1 and associates with actin filaments and stress fibers in activated platelets and endothelial cells.
11101506 2000 The interaction of titin and alpha-actinin is controlled by a phospholipid-regulated intramolecular pseudoligand mechanism.
10926853 2000 Fine mapping of the alpha-actinin binding site within cysteine-rich protein.
10833443 2000 ENH, containing PDZ and LIM domains, heart/skeletal muscle-specific protein, associates with cytoskeletal proteins through the PDZ domain.
10801846 2000 Cardiac phospholipase D2 localizes to sarcolemmal membranes and is inhibited by alpha-actinin in an ADP-ribosylation factor-reversible manner.
10793131 2000 Identification of a novel member of the chloride intracellular channel gene family (CLIC5) that associates with the actin cytoskeleton of placental microvilli.
10470015 1999 Functional dissection of nebulette demonstrates actin binding of nebulin-like repeats and Z-line targeting of SH3 and linker domains.
10369880 1999 Myotilin, a novel sarcomeric protein with two Ig-like domains, is encoded by a candidate gene for limb-girdle muscular dystrophy.
10224105 1999 An alpha-actinin binding site of zyxin is essential for subcellular zyxin localization and alpha-actinin recruitment.
10198040 1999 Protein interactions with the glucose transporter binding protein GLUT1CBP that provide a link between GLUT1 and the cytoskeleton.
10029173 1999 A fragment of alpha-actinin promotes monocyte/macrophage maturation in vitro.
9599291 1998 Integrins and development: how might these receptors regulate differentiation of the lens.
9030526 1997 Interaction of PKN with alpha-actinin.
8713105 1996 Identification of the 70kD heat shock cognate protein (Hsc70) and alpha-actinin-1 as novel phosphotyrosine-containing proteins in T lymphocytes.
8125298 1994 Oligo-capping: a simple method to replace the cap structure of eukaryotic mRNAs with oligoribonucleotides.
7983147 1994 Modulation of alpha-actinin levels affects cell motility and confers tumorigenicity on 3T3 cells.
7829101 1994 Isolation of novel and known genes from a human fetal cochlear cDNA library using subtractive hybridization and differential screening.
7790378 1995 Interaction of alpha-actinin with the cadherin/catenin cell-cell adhesion complex via alpha-catenin.
7750553 1995 Identification of the cytoskeletal protein alpha-actinin as a platelet thrombospondin-binding protein.
2780298 1989 The cDNA sequence of a human placental alpha-actinin.
2349951 1990 Cloning and chromosomal localization of the human cytoskeletal alpha-actinin gene reveals linkage to the beta-spectrin gene.
2169343 1990 Expression of human alpha-actinin in human hepatocellular carcinoma.
1720388 1991 Transfection of chicken skeletal muscle alpha-actinin cDNA into nonmuscle and myogenic cells: dimerization is not essential for alpha-actinin to bind to microfilaments.
1629252 1992 Alpha-actinin and vinculin in human neutrophils: reorganization during adhesion and relation to the actin network.