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Lipopolysaccharide
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Structure of a lipopolysaccharide
Lipopolysaccharides (LPS), also known as lipoglycans and endotoxins, are large molecules consisting of a lipid and a polysaccharide composed of O-antigen, outer core and inner core joined by a covalent bond; they are found in the outer membrane of Gram-negative bacteria, and elicit strong immune responses in animals.
The term lipooligosaccharide ("LOS") is used to refer to a low-molecular-weight form of bacterial lipopolysaccharides.
Contents
[hide]
1 Discovery
2 Functions in bacteria
3 Composition
3.1 O-antigen
3.2 Core
3.3 Lipid A
4 Lipooligosaccharides
5 LPS modifications
6 Biosynthesis and transport
7 Biological effects on hosts infected with gram-negative bacteria
7.1 Immune response
7.2 Effect of variability on immune response
8 Health effects
8.1 Endotoxemia
8.2 Auto-immune disease
8.3 Link to obesity
9 Laboratory research and biotechnology production systems
10 See also
11 References
12 External links
Discovery[edit]
The toxic activity of LPS was first discovered and termed "endotoxin" by Richard Friedrich Johannes Pfeiffer, who distinguished between exotoxins, which he classified as a toxin that is released by bacteria into the surrounding environment, and endotoxins, which he considered to be a toxin kept "within" the bacterial cell and released only after destruction of the bacterial cell wall.[1]:84 Subsequent work showed that release of LPS from gram negative microbes does not necessarily require the destruction of the bacterial cell wall, but rather, LPS is secreted as part of the normal physiological activity of membrane vesicle trafficking in the form of bacterial outer membrane vesicles (OMVs), which may also contain other virulence factors and proteins.[2]
Today, the term 'endotoxin' is mostly used synonymously with LPS,[3] although there are a few endotoxins that are not related to LPS, such as the so-called delta endotoxin proteins secreted by Bacillus thuringiensis.
Functions in bacteria[edit]
LPS is the major component of the outer membrane of Gram-negative bacteria, contributing greatly to the structural integrity of the bacteria, and protecting the membrane from certain kinds of chemical attack. LPS also increases the negative charge of the cell membrane and helps stabilize the overall membrane structure. It is of crucial importance to Gram-negative bacteria, which die if it is mutated or removed. LPS induces a strong response from normal animal immune systems. It has also been implicated in non-pathogenic aspects of bacterial ecology, including surface adhesion, bacteriophage sensitivity, and interactions with predators such as amoebae.
LPS is required for the proper conformation of Omptin activity; however, smooth LPS will sterically hinder omptins.
Composition[edit]
The saccharolipid Kdo2-Lipid A. Kdo residues in red (core), glucosamine residues in blue, acyl chains in black and phosphate groups in green.
It comprises three parts:
O antigen (or O polysaccharide)
Core oligosaccharide
Lipid A
O-antigen[edit]
A repetitive glycan polymer contained within an LPS is referred to as the O antigen, O polysaccharide, or O side-chain of the bacteria. The O antigen is attached to the core oligosaccharide, and comprises the outermost domain of the LPS molecule. The composition of the O chain varies from strain to strain. For example, there are over 160 different O antigen structures produced by different E. coli strains.[4] The presence or absence of O chains determines whether the LPS is considered rough or smooth. Full-length O-chains would render the LPS smooth, whereas the absence or reduction of O-chains would make the LPS rough.[5] Bacteria with rough LPS usually have more penetrable cell membranes to hydrophobic antibiotics, since a rough LPS is more hydrophobic.[6] O antigen is exposed on the very outer surface of the bacterial cell, and, as a consequence, is a target for recognition by host antibodies.
Core[edit]
Main article: Core oligosaccharide
The Core domain always contains an oligosaccharide component that attaches directly to lipid A and commonly contains sugars such as heptose and 3-Deoxy-D-manno-oct-2-ulosonic acid (also known as KDO, keto-deoxyoctulosonate).[7] The LPS Cores of many bacteria also contain non-carbohydrate components, such as phosphate, amino acids, and ethanolamine substituents.
Lipid A[edit]
Main article: Lipid A
Lipid A is, in normal circumstances, a phosphorylated glucosamine disaccharide decorated with multiple fatty acids. These hydrophobic fatty acid chains anchor the LPS into the bacterial membrane, and the rest of the LPS projects from the cell surface. The lipid A domain is responsible for much of the toxicity of Gram-negative bacteria. When bacterial cells are lysed by the immune system, fragments of membrane containing lipid A are released into the circulation, causing fever, diarrhea, and possible fatal endotoxic shock (also called septic shock). The Lipid A moiety is a very conserved component of the LPS.[8]
Lipooligosaccharides[edit]
Lipooligosaccharides (LOS) are glycolipids found in the outer membrane of some types of Gram-negative bacteria, such as Neisseria spp. and Haemophilus spp. The term is synonymous with the low molecular weight form of bacterial LPS.[9] LOS plays a central role in maintaining the integrity and functionality of the outer membrane of the Gram negative cell envelope. Lipooligosaccharides play an important role in the pathogenesis of certain bacterial infections because they are capable of acting as immunostimulators and immunomodulators.[9] Furthermore, LOS molecules are responsible for the ability of some bacterial strains to display molecular mimicry and antigenic diversity, aiding in the evasion of host immune defenses and thus contributing to the virulence of these bacterial strains.
Chemically, lipooligosaccharides lack O-antigens and possess only a lipid A-based outer membrane-anchoring moiety, and an oligosaccharide core.[10] In the case of Neisseria meningitidis, the lipid A portion of the molecule has a symmetrical structure and the inner core is composed of 3-deoxy-D-manno-2-octulosonic acid (KDO) and heptose (Hep) moieties. The outer core oligosaccharide chain varies depending on the bacterial strain.[9][10] The term lipooligosaccharide is used to refer to the low molecular weight form of bacterial lipopolysaccharides, which can be categorized into two forms: the high molecular weight (Mr, or smooth) form possesses a high molecular weight, repeating polysaccharide O-chain, while the low molecular weight (low-Mr or rough) form, lacks the O-chain but possesses a short oligosaccharide in its place.[9]
LPS modifications[edit]
The making of LPS can be modified in order to present a specific sugar structure. Those can be recognised by either other LPS (which enables to inhibit LPS toxins) or glycosyltransferases that use those sugar structure to add more specific sugars. It has recently been shown that a specific enzyme in the intestine (alkaline phosphatase) can detoxify LPS by removing the two phosphate groups found on LPS carbohydrates.[11] This may function as an adaptive mechanism to help the host manage potentially toxic effects of gram-negative bacteria normally found in the small intestine. A different enzyme may detoxify LPS when it enters, or is produced in, animal tissues. Neutrophils, macrophages, and dendritic cells produce a lipase, acyloxyacyl hydrolase (AOAH), that inactivates LPS by removing the two secondary acyl chains from lipid A. If they are given LPS parenterally, mice that lack AOAH develop high titers of non-specific antibodies, develop prolonged hepatomegaly, and experience prolonged endotoxin tolerance. LPS inactivation may be required for animals to restore homeostasis after parenteral LPS exposure.[12]
Biosynthesis and transport[edit]
LPS Final Assembly: O-antigen subunits are translocated across the inner membrane (by Wzx) where they are polymerized (by Wzy, chain length determined by Wzz) and ligated (by WaaL) on to complete Core-Lipid A molecules (which were translocated by MsbA).[13]
LPS Transport: Completed LPS molecules are transported across the periplasm and outer membrane by the proteins LptA, B, C, D, E, F, and G[14]
Biological effects on hosts infected with gram-negative bacteria[edit]
Immune response[edit]
LPS acts as the prototypical endotoxin because it binds the CD14/TLR4/MD2 receptor complex in many cell types, but especially in monocytes, dendritic cells, macrophages and B cells, which promotes the secretion of pro-inflammatory cytokines, nitric oxide, and eicosanoids.[15]
As part of the cellular stress response, superoxide is one of the major reactive oxygen species induced by LPS in various cell types that express TLR (toll-like receptor).
LPS is also an exogenous pyrogen (fever-inducing substance).
Being of crucial importance to Gram-negative bacteria, these molecules make candidate targets for new antimicrobial agents.
Some researchers doubt reports of generalized toxic effects attributed to all lipopolysaccharides, in particular, for cyanobacteria.[16]
LPS function has been under experimental research for several years due to its role in activating many transcription factors. LPS also produces many types of mediators involved in septic shock. Humans are much more sensitive to LPS than other animals (e.g., mice). A dose of 1 µg/kg induces shock in humans, but mice will tolerate a dose up to a thousand times higher.[17] This may relate to differences in the level of circulating natural antibodies between the two species.[18][19] Said et al. showed that LPS causes an IL-10-dependent inhibition of CD4 T-cell expansion and function by up-regulating PD-1 levels on monocytes which leads to IL-10 production by monocytes after binding of PD-1 by PD-L1.[20]
Endotoxins are in large part responsible for the dramatic clinical manifestations of infections with pathogenic Gram-negative bacteria, such as Neisseria meningitidis, the pathogens that causes meningococcal disease, including meningococcemia, Waterhouse–Friderichsen syndrome, and meningitis.
Bruce Beutler was awarded a portion of the 2011 Nobel Prize in Physiology or Medicine for his work demonstrating that TLR4 is the LPS receptor.[21][22]
Portions of the LPS from several bacterial strains have been shown to be chemically similar to human host cell surface molecules; the ability of some bacteria to present molecules on their surface which are chemically identical or similar to the surface molecules of some types of host cells is termed molecular mimicry.[23] For example, in Neisseria meningitidis L2,3,5,7,9, the terminal tetrasaccharide portion of the oligosaccharide (lacto-N-neotetraose) is the same tetrasaccharide as that found in paragloboside, a precursor for ABH glycolipid antigens found on human erythrocytes.[9] In another example, the terminal trisaccharide portion (lactotriaose) of the oligosaccharide from pathogenic Neisseria spp. LOS is also found in lactoneoseries glycosphingolipids from human cells.[9] Most meningococci from groups B and C, as well as gonococci, have been shown to have this trisaccharide as part of their LOS structure.[9] The presence of these human cell surface 'mimics' may, in addition to acting as a 'camouflage' from the immune system, play a role in the abolishment of immune tolerance when infecting hosts with certain human leukocyte antigen (HLA) genotypes, such as HLA-B35.[9]
Effect of variability on immune response[edit]
Toll-like receptors of the innate immune system recognize LPS and trigger an immune response.
O-antigens (the outer carbohydrates) are the most variable portion of the LPS molecule, imparting the antigenic specificity. In contrast, lipid A is the most conserved part. However, lipid A composition also may vary (e.g., in number and nature of acyl chains even within or between genera). Some of these variations may impart antagonistic properties to these LPS. For example, Rhodobacter sphaeroides diphosphoryl lipid A (RsDPLA) is a potent antagonist of LPS in human cells, but is an agonist in hamster and equine cells.[citation needed]
It has been speculated that conical Lipid A (e.g., from E. coli) are more agonistic, less conical lipid A like those of Porphyromonas gingivalis may activate a different signal (TLR2 instead of TLR4), and completely cylindrical lipid A like that of Rhodobacter sphaeroides is antagonistic to TLRs.[24][25]
LPS gene clusters are highly variable between different strains, subspecies, species of bacterial pathogens of plants and animals.[26][27]
Normal human blood serum contains anti-LOS antibodies that are bactericidal and patients that have infections caused by serotypically distinct strains possess anti-LOS antibodies that differ in their specificity compared with normal serum.[28] These differences in humoral immune response to different LOS types can be attributed to the structure of the LOS molecule, primarily within the structure of the oligosaccharide portion of the LOS molecule.[28] In Neisseria gonorrhoeae it has been demonstrated that the antigenicity of LOS molecules can change during an infection due to the ability of these bacteria to synthesize more than one type of LOS,[28] a characteristic known as phase variation. Additionally, Neisseria gonorrhoeae, as well as Neisseria meningitidis and Haemophilus influenzae,[9] are capable of further modifying their LOS in vitro, for example through sialylation (modification with sialic acid residues), and as a result are able to increase their resistance to complement-mediated killing [28] or even down-regulate complement activation[9] or evade the effects of bactericidal antibodies.[9] Sialylation may also contribute to hindered neutrophil attachment and phagocytosis by immune system cells as well as a reduced oxidative burst.[9] Haemophilus somnus, a pathogen of cattle, has also been shown to display LOS phase variation, a characteristic which may help in the evasion of bovine host immune defenses.[29] Taken together, these observations suggest that variations in bacterial surface molecules such as LOS can help the pathogen evade both the humoral (antibody and complement-mediated) and the cell-mediated (killing by neutrophils, for example) host immune defenses.
Health effects[edit]
Endotoxemia[edit]
The presence of endotoxins in the blood is called endotoxemia. It can lead to septic shock, if the immune response is severely pronounced.[30]
Moreover, endotoxemia of intestinal origin, especially, at the host-pathogen interface, is considered to be an important factor in the development of alcoholic hepatitis,[31] which is likely to develop on the basis of the small bowel bacterial overgrowth syndrome and an increased intestinal permeability.[32]
Lipid A may cause uncontrolled activation of mammalian immune systems with production of inflammatory mediators that may lead to septic shock.[10] This inflammatory reaction is mediated by Toll-like receptor 4 which is responsible for immune system cell activation.[10] Damage to the endothelial layer of blood vessels caused by these inflammatory mediators can lead to capillary leak syndrome, dilation of blood vessels and a decrease in cardiac function and can lead to septic shock.[33] Pronounced complement activation can also be observed later in the course as the bacteria multiply in the blood.[33] High bacterial proliferation triggering destructive endothelial damage can also lead to disseminated intravascular coagulation (DIC) with loss of function of certain internal organs such as the kidneys, adrenal glands and lungs due to compromised blood supply. The skin can show the effects of vascular damage often coupled with depletion of coagulation factors in the form of petechiae, purpura and ecchymoses. The limbs can also be affected, sometimes with devastating consequences such as the development of gangrene, requiring subsequent amputation.[33] Loss of function of the adrenal glands can cause adrenal insufficiency and additional hemorrhage into the adrenals causes Waterhouse-Friderichsen syndrome, both of which can be life-threatening. It has also been reported that gonococcal LOS can cause damage to human fallopian tubes.[28]
Auto-immune disease[edit]
The molecular mimicry of some LOS molecules is thought to cause autoimmune-based host responses, such as flareups of multiple sclerosis.[9][23] Other examples of bacterial mimicry of host structures via LOS are found with the bacteria Helicobacter pylori and Campylobacter jejuni, organisms which cause gastrointestinal disease in humans, and Haemophilus ducreyi which causes chancroid. Certain C. jejuni LPS serotypes (attributed to certain tetra- and pentasaccharide moieties of the core oligosaccharide) have also been implicated with Guillain–Barré syndrome and a variant of Guillain–Barré called Miller-Fisher syndrome.[9]
Link to obesity[edit]
Epidemiological studies have previously shown that increased endotoxin load, which can be a result of increased populations of endotoxin producing bacteria in the intestinal tract, is associated with certain obesity-related patient groups.[34][35][36] Other studies have shown that purified endotoxin from Escherichia coli can induce obesity and insulin-resistance when injected into germ-free mouse models.[37] A more recent study has uncovered a potentially contributing role for Enterobacter cloacae B29 toward obesity and insulin resistance in a human patient.[38] The presumed mechanism for the association of endotoxin with obesity is that endotoxin induces an inflammation-mediated pathway accounting for the observed obesity and insulin resistance.[37]There is a correlation between plasma LPS and insulin resistance.[39]
Bacterial genera associated with endotoxin-related obesity effects: Escherichia, Enterobacter
Laboratory research and biotechnology production systems[edit]
Lipopolysaccharides are frequent contaminants in plasmid DNA prepared from bacteria or proteins expressed from bacteria, and must be removed from the DNA or protein to avoid contaminating experiments and to avoid toxicity of products manufactured using industrial fermentation.[40]
Also, ovalbumin is frequently contaminated with endotoxins. Ovalbumin is one of the extensively studied proteins in animal models and also an established model allergen for airway hyper-responsiveness (AHR). Commercially available ovalbumin that is contaminated with LPS can fully activate endothelial cells in an in-vitro assay of the first step of inflammation, and it falsifies research results, as it does not accurately reflect the effect of sole protein antigen on animal physiology.[citation needed]
In pharmaceutical production, it is necessary to remove all traces of endotoxin from drug product containers, as even small amounts of endotoxin will cause illness in humans. A depyrogenation oven is used for this purpose. Temperatures in excess of 300 °C are required to break down this substance. A defined endotoxin reduction rate is a correlation between time and temperature. Based on primary packaging material as syringes or vials, a glass temperature of 250 °C and a holding time of 30 minutes is typical to achieve a reduction of endotoxin levels by a factor of 1000.[41]
The standard assay for detecting presence of endotoxin is the Limulus Amebocyte Lysate (LAL) assay, utilizing blood from the Horseshoe crab (Limulus polyphemus).[42] Very low levels of LPS can cause coagulation of the limulus lysate due to a powerful amplification through an enzymatic cascade. However, due to the dwindling population of horseshoe crabs, and the fact that there are factors that interfere with the LAL assay, efforts have been made to develop alternative assays, with the most promising ones being ELISA tests using a recombinant version of a protein in the LAL assay, Factor C.[43]
See also[edit]
Bioaerosol
Depyrogenation
Exotoxin
Host-pathogen interface
Membrane vesicle trafficking
Mucopolysaccharide
Nesfatin-1
Schwartzman reaction
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Jump up ^ Iwanaga S (2007). "Biochemical principle of Limulus test for detecting bacterial endotoxins". Proc. Jpn. Acad., Ser. B, Phys. Biol. Sci. 83 (4): 110–9. PMC 3756735  . PMID 24019589. doi:10.2183/pjab.83.110.
Jump up ^ Ding JL, Ho B (2001). "A new era in pyrogen testing" (PDF). Trends Biotechnol. 19 (8): 277–81. PMID 11451451. doi:10.1016/s0167-7799(01)01694-8.
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Subliminal stimuli
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This article needs more medical references for verification or relies too heavily on primary sources. Please review the contents of the article and add the appropriate references if you can. Unsourced or poorly sourced material may be challenged and removed. (August 2015)
Subliminal stimuli (/sʌbˈlɪmɪnəl/; literally "below threshold"), contrary to supraliminal stimuli or "above threshold", are any sensory stimuli below an individual's threshold for conscious perception.[1] A recent review of functional magnetic resonance imaging (fMRI) studies shows that subliminal stimuli activate specific regions of the brain despite participants being unaware.[2] Visual stimuli may be quickly flashed before an individual can process them, or flashed and then masked, thereby interrupting the processing. Audio stimuli may be played below audible volumes or masked by other stimuli.
Contents
[hide]
1 Effectiveness
2 Method
2.1 Objective threshold
2.2 Subjective threshold
2.3 Direct and indirect measures
3 Visual stimuli
3.1 Images
3.2 Emotion eliciting stimuli
3.3 Simple geometric stimuli
3.4 Word and non-word stimuli
3.5 Masking visual stimuli
4 Auditory stimuli
4.1 Auditory masking
4.2 Self-help audio recordings
5 Consumption and television
6 See also
7 References
8 Further reading
Effectiveness[edit]
Applications of subliminal stimuli often base themselves on the persuasiveness of the message. Importantly, research on action priming has shown that subliminal stimuli can trigger only actions a receiver of the message plans to perform anyway; however, consensus of this finding remains unsubstantiated by other research. Most actions can be triggered subliminally only if the person already has the potential to perform a specific action.[3] The following sections have more information on specific studies which investigate the effectiveness of subliminal stimuli.[4][5]
Method[edit]
The threshold in subliminal stimuli research is the level at which the participant is not aware of the stimulus being presented.[6] Researchers determine a threshold for the stimulus that will be used as the subliminal stimulus. That subliminal stimulus is then presented during the study at some point and measures are taken to determine the effects of the stimulus. The way in which studies operationally define thresholds depends on the methods of the particular article. The methodology of the research also varies by the type of subliminal stimulus (auditory or visual) and the dependent variables they measure.
Objective threshold[edit]
The objective threshold is found using a forced choice procedure, in which participants must choose which stimulus they saw from options given to them.[6] Participants are flashed a stimulus (e.g. the word "orange") and then given a few choices and asked which one they saw. Participants must choose an answer in this design. The objective threshold is obtained when participants are at chance level of performance in this task.[6] The length of presentation that causes chance performance on the forced choice task is used later in the study for the subliminal stimuli.
Subjective threshold[edit]
The subjective threshold is determined by when the participant reports that his or her performance on the forced choice procedure approximates chance.[6] The subjective threshold is 30 to 50 ms slower than the objective threshold, demonstrating that participants are able to detect the stimuli is present sooner than their perceived accuracy ratings would indicate.[6] In other words, stimuli presented at a subjective threshold have a longer presentation time than those presented at an objective threshold. When using the objective threshold, primes neither facilitated nor inhibited the recognition of a color.[6] However, the longer the duration of the priming stimuli, the greater effect it had on subsequent responding.[6] These findings indicate that the results of some studies may be due to their definition of below threshold.
Direct and indirect measures[edit]
Perception without awareness can be demonstrated through the comparison of direct and indirect measures of perception.[7] Direct measures use responses to task definitions as per the explicit instructions given to the subjects. Indirect measures use responses that are not a part of the task definition given to subjects.[7] Both direct and indirect measures are displayed under comparable conditions except for the direct or indirect instruction. For example, in a typical Stroop Task, subjects are given the task of naming the color of a patch of ink. A direct measure is 'accuracy'—true to the instructions given to the participants. The popular indirect measure used in the same task is 'response time'—subjects are not told that they are being measured for response times.
Similarly, a direct effect is the effect of a task stimulus on the instructed response to that stimulus, usually measured as accuracy. An indirect effect is an uninstructed effect of the task stimulus on behavior, sometimes measured by including an irrelevant or distracting component in the task stimulus and measuring its effect on accuracy.[8] These effects are then compared on their relative sensitivity: an indirect effect that is greater than direct effect indicates existence of unconscious cognition.[7][8]
Visual stimuli[edit]
In order to study the effects of subliminal stimuli, researchers will often prime the participants with specific visual stimuli, often images, and determine if those stimuli elicit different responses.[9][10] Subliminal stimuli have mostly been studied in the context of emotion, in particular, researchers have focused a lot of attention to the perception of faces and how subliminal presentation to different facial expression affects emotion.[11][12][13][14] Visual subliminal stimuli has also been used to study emotion eliciting stimuli and simple geometric stimuli.[15][16][17][18] A significant amount of research has been produced throughout the years to demonstrate the effects of subliminal visual stimuli.[19]
Images[edit]
Attitudes can develop without being aware of its antecedents.[9] Individuals viewed slides of people performing familiar daily activities after being exposed to either an emotionally positive scene, such as a romantic couple or kittens, or an emotionally negative scene, such as a werewolf or a dead body between each slide and the next. After exposure from something which the individuals consciously perceived as a flash of light, the participants gave more positive personality traits to those people whose slides were associated with an emotionally positive scene and vice versa. Despite the statistical difference, the subliminal messages [20] had less of an impact on judgment than the slide's inherent level of physical attractiveness.[9]
Individuals show right amygdala activity in response to subliminal fear, and greater left amygdala response to supraliminal fear.[10] People were exposed to a subliminal image flashed for 16.7 milliseconds that could signal a potential threat and again with a supraliminal image flashed for half a second. Furthermore, supraliminal fear showed more sustained cortical activity, suggesting that subliminal fear may not entail conscious surveillance while supraliminal fear entails higher-order processing.[10]
Emotion eliciting stimuli[edit]
A subliminal sexual stimulus has a different effect on men compared to women.[15] Men and women were subliminally exposed to either a sexual or a neutral picture, and their sexual arousal was recorded. Researchers examined the accessibility of sex-related thoughts after following the same procedure with either a pictorial judgment task or lexical decision task. The results revealed that the subliminal sexual stimuli did not have an effect on men, but for women, lower levels of sexual arousal were reported.[15] However, in conditions related to accessibility of sex-related thoughts, the subliminal sexual stimuli led to higher accessibility for both men and women.[15]
Subliminal stimuli can elicit significant emotional changes, but these changes are not valuable for a therapeutic effect.[16] Spider-fearful and non-fearful undergraduates experienced either a positive, negative, or neutral subliminal prime followed immediately by a picture of a spider or a snake. Using visual analogue scales, the participants rated the affective quality of the picture. No evidence was found to support that the unpleasantness of the pictures can be modulated by subliminal priming.[16] In fact, the non-fearful participants rated the spiders as more frightening after being primed with a negative stimuli, however, for the fearful participants, this effect was not found.[16]
Simple geometric stimuli[edit]
Laboratory research on unconscious perception often employs simple stimuli (e.g., geometric shapes or colors) in which visibility is controlled by visual masking.[17] Masked stimuli are then used to prime the processing of subsequently presented target stimuli. For instance, in the Response Priming paradigm, participants have to respond to a target stimulus (e.g., by identifying whether it is a diamond or a square) which is immediately preceded by a masked priming stimulus (also a diamond or a square). The prime has large effects on responses to the target; it speeds responses when it is consistent with the target, and slows responses when it is inconsistent. Response priming effects can be dissociated from visual awareness of the prime, such as when prime identification performance is at chance, or when priming effects increase despite decreases in prime visibility.[17]
The presentation of geometric figures as subliminal stimuli can result in below threshold discriminations.[18] The geometric figures were presented on slides of a tachistoscope followed by a supraliminal shock for a given slide every time it appeared. The shock was administered after a five-second interval. Electrical skin changes of the participants that occurred before the reinforcement (shock) or non-reinforcement were recorded. The findings indicate that the proportion of electrical skin changes that occurred following subliminal visual stimuli was significantly greater than expected.[18] In contrast, the proportion of electrical skin changes that occurred in response to the stimuli which were not reinforced was significantly less. As a whole, participants were able to make below threshold discriminations.[18]
Word and non-word stimuli[edit]
Another form of visual stimuli are words and non-words. In a set of experiments, words and non-word were used as subliminal primes. Primes that work best as subliminal stimuli are words that have been classified several times before they are used as primes. Word primes can also be made from parts of practiced words to create new words. In this case, the actual word used as a prime can have the opposite meaning of the words it came from (its “parents”), but it will still prime for the meaning of the parent words. Non-words created from previously practiced stimuli have a similar effect, even when they are unpronounceable (e.g., made of all consonants). These primes generally only increase response times for later stimuli for a very short period of time (milliseconds).[21]
Masking visual stimuli[edit]
Visual stimuli are often masked by forward and backward masks so that they can be displayed for longer periods of time, but without the subject being able to tell what the prime is. A forward mask displayed before the prime for a short period of time and usually a backward mask will follow the prime. This prevents recognition of the prime by the subject.[22]
Auditory stimuli[edit]
Auditory masking[edit]
One method for creating subliminal auditory stimuli is called masking. This method involves hiding the target auditory stimulus in some way. Auditory subliminal stimuli are shown to have some effect on the participant, but not a large one.[22] For example, one study used other speechlike sounds to cover up the target words.[22] The study found evidence of priming in the absence of awareness of the stimuli. But the effects of these subliminal stimuli were only seen in one of the outcome measures of priming, while the effects of conscious stimuli were seen in multiple outcome measures.[22] However, the empirical evidence for the assumption of an impact of auditory subliminal stimuli on human behavior remains weak: In an experimental study on the influence of subliminal target words (embedded into a music track) on choice behavior for a drink,[23] authors found no evidence for a manipulative effect.
Self-help audio recordings[edit]
A study investigated the effects on self-concept of Rational Emotive Behavior Therapy and auditory subliminal stimulation (separately and in combination) on 141 undergraduate students with self-concept problems. They were randomly assigned to one of four groups receiving either Rational-Emotive Therapy, subliminal stimulation, both, or a placebo treatment. Rational-Emotive Therapy significantly improved scores on all the dependent measures (cognition, self-concept, self-esteem, anxiety), except for behavior. Results for the subliminal stimulation group were similar to those of the placebo treatment except for a significant self-concept improvement and a decline in self-concept related irrational cognitions. The combined treatment yielded results similar to those of Rational-Emotive Therapy, with tentative indications of continued improvement in irrational cognitions and self-concept from posttest to follow-up.[24]
Another study investigated the effects of self-help tapes on self-esteem and memory. Volunteers who wanted to improve their self-esteem or memory were recruited and completed several self-esteem and memory tests before being given a self-help tape. Subjects were given a self-esteem audio tape or a memory audio tape, but half of the tapes were mislabeled and half were correctly labeled. After listening to the tapes daily for five weeks, subjects came back and repeated the self-esteem and memory tests. There was no significant change from the first set of testing to the second, although subjects believed that their self-esteem or memory improved based on which tape they believed they had, even when they had a mislabeled tape (those who had tapes labeled as self-esteem tapes felt their self-esteem had increased and the same with memory). This effect is often referred to as a placebo.[25][26] There are multiple other studies on subliminal self-help with different measures and have given similar results.[27][28]
Consumption and television[edit]
Some studies have looked at the efficacy of subliminal messaging in television. Subliminal messages produce only one-tenth of the effects of detected messages and the findings related the effects of subliminal messaging were relatively ambiguous.[29] Also, participants’ ratings of positive response to commercials are not affected by subliminal messages in the commercials.[29]
Karremans suggests that subliminal messages have an effect when the messages are goal-relevant.[30] Subliminally priming a brand name of a soft drink (Lipton Ice) made those who were thirsty want the Lipton Ice. However, those who were not thirsty were not influenced by the subliminal messages.[30] Karremans did a study assessing whether subliminal priming of a brand name of a drink would affect a person's choice of drink, and whether this effect is caused by the individual's feelings of being thirsty.[30] In another study, participant's ratings of thirst were higher after viewing an episode of The Simpsons that contained single frames of the word "thirsty" or of a picture of a Coca-Cola can.[31] Some studies have shown greater effects of subliminal messaging with as high as 80% of participants showing a preference for a particular rum when subliminally primed by the name placed in an ad backward.[32]
Many authors have continued to argue for the effectiveness of subliminal cues in changing consumption behavior, citing environmental cues as a main culprit of behavior change.[33] Authors who support this line of reasoning cite findings such as the research that showed slow-paced music in a supermarket was associated with more sales and customers moving at a slower pace.[34] Findings such as these support the notion that external cues can affect behavior, although the stimulus may not fit into a strict definition of subliminal stimuli because although the music may not be attended to or consciously affecting the customers, they are certainly able to perceive it.
Subliminal messaging is prohibited in advertising in the United Kingdom.[35]
See also[edit]
Instances of subliminal messages
Neuroethics
Peripheral vision horizon display (PVHD)
Pre-attentive processing
Priming (psychology)
Response priming
Unconscious thought theory § Criticism of UTT
They Live
References[edit]
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Jump up ^ Brooks, S.J.; Savov V; Allzén E; Benedict C; Fredriksson R; Schiöth HB. (February 2012). "Exposure to subliminal arousing stimuli induces robust activation in the amygdala, hippocampus, anterior cingulate, insular cortex and primary visual cortex: a systematic meta-analysis of fMRI studies.". NeuroImage. 59 (3): 2962–2973. PMID 22001789. doi:10.1016/j.neuroimage.2011.09.077.
Jump up ^ Schlaghecken, F.; Eimer, M. (2004). "Subliminal stimuli can bias 'free' choices between response alternatives". Psychonomic Bulletin & Review. 11: 463–468. doi:10.3758/bf03196596.
Jump up ^ van Gaal, S; de Lange, FP; Cohen, MX (2012). "The role of consciousness in cognitive control and decision making". Front Hum Neurosci. 6: 121. PMC 3345871  . PMID 22586386. doi:10.3389/fnhum.2012.00121.
Jump up ^ Zeman, A. (2001). "Consciousness". Brain. 124 (Pt 7): 1263–1289. PMID 11408323. doi:10.1093/brain/124.7.1263.
^ Jump up to: a b c d e f g Chessman, Jim; Merikle, Philip M. (1984). "Priming with and without awareness". Perception and Psychophysics. 36 (4): 387–395. doi:10.3758/bf03202793.
^ Jump up to: a b c Reingold, Eyal M.; Merikle, Philip M. (1988). "Direct and indirect measures to study perceptions without awareness". Perception and Psychophysics. 44 (6): 563–575. doi:10.3758/bf03207490.
^ Jump up to: a b Greenwald, Anthony G.; Klinger, Mark R.; Schuh, Eric S. (1995). "Activation by marginally Perceptible ("Subliminal") Stimuli: Dissociation of Unconscious From Conscious Cognition". Journal of Experimental Psychology. 124 (1): 22–42. doi:10.1037/0096-3445.124.1.22.
^ Jump up to: a b c Krosnick, J. A.; Betz, A. L.; Jussim, L. J.; Lynn, A. R. (1992). "Subliminal Conditioning of Attitudes". Personality and Social Psychology Bulletin. 18 (2): 152–162. doi:10.1177/0146167292182006.
^ Jump up to: a b c Williams, L. M.; Liddell, B. J.; Kemp, A. H.; Bryant, R. A.; Meares, R. A.; Peduto, A. S.; Gordon, E. (2006). "Amygdala–prefrontal dissociation of subliminal and supraliminal fear". Human Brain Mapping. 27 (8): 652–661. PMID 16281289. doi:10.1002/hbm.20208.
Jump up ^ Baldwin, M.; Carrell, D. F.; Lopez, D. F. (1990). "Priming relationship schemas: My advisor and the pope are watching me from the back of my mind". Journal of Experimental Social Psychology. 26 (5): 435–454. doi:10.1016/0022-1031(90)90068-W.
Jump up ^ Lee, Su Young; Kang, Jee In; Lee, Eun; Namkoong, Kee; An, Suk Kyoon (February 2011). "Differential priming effect for subliminal fear and disgust facial expressions". Attention, Perception, & Psychophysics. 2. 73: 473–481. doi:10.3758/s13414-010-0032-3.
Jump up ^ Ibáñez, Agustin; Hurtado, Esteban; Lobos, Alejandro; Escobar, Josefina; Trujillo, Natalia; Baez, Sandra; Huepe, David; Manes, Facundo; Decety, Jean (29 June 2011). "Subliminal presentation of other faces (but not own face) primes behavioral and evoked cortical processing of empathy for pain". Brain Research. 1398: 72–85. PMID 21624566. doi:10.1016/j.brainres.2011.05.014.
Jump up ^ Yang, Zixu; Tong, Eddie M. W. (2010). "The Effects of Subliminal Anger and Sadness Primes on Agency Appraisals". Emotion. 10 (6): 915–922. doi:10.1037/a0020306.
^ Jump up to: a b c d Gillath, Omri; Mikulincer, Mario; Birnbaum, Gurit E.; Shaver, Phillip R. (May 2007). "Does subliminal exposure to sexual stimuli have the same effects on men women?". Journal of Sex Research. 2. 44: 111–121. doi:10.1080/00224490701263579.
^ Jump up to: a b c d Mayer, Birgit; Merckelbach, Harald (10 December 1998). "Do subliminal priming effects on emotion have clinical potential?". Anxiety, Stress, & Coping. 12: 217–229. doi:10.1080/10615809908248330.
^ Jump up to: a b c Vorberg, D.; Mattler, U.; Heinecke, A.; Schmidt, T.; Schwarzbach, J. (2003). "Different time courses for visual perception and action priming". Proceedings of the National Academy of Sciences USA. 100: 6275–6280. doi:10.1073/pnas.0931489100.
^ Jump up to: a b c d Taylor, Frank W. R. (March 1953). "The discrimination of subliminal visual stimuli". Canadian Journal of Experimental Psychology. 1. 7: 12–20. doi:10.1037/h0083570.
Jump up ^ Kouider, S; Dehaene, S (May 2007). "Levels of processing during non-conscious perception: a critical review of visual masking.". Philos Trans R Soc Lond B Biol Sci. 362: 857–75. PMC 2430002  . PMID 17403642. doi:10.1098/rstb.2007.2093.
Jump up ^ www.monyms.ir
Jump up ^ Abrams, R. L.; Greenwald, A. G. (2000). "Parts outweigh the whole (word) in unconscious analysis of meaning". Psychological Science. 11: 118–124. doi:10.1111/1467-9280.00226.
^ Jump up to: a b c d Breitmeyer, B.G.; Ogmen, H. (2007). "Visual Masking". Scholarpedia. 2 (7): 3330. doi:10.4249/scholarpedia.3330.
Jump up ^ Egermann, Hauke; Kopiez, Reinhard; Reuter, Christoph (2007). "Is there an effect of subliminal messages in music on choice behavior?". Journal of Articles in Support of the Null Hypothesis go. 4 (2): 29–64.
Jump up ^ Comparison of the effects of auditory subliminal stimulation and rational-emotive therapy, separately and combined, on self-concept. Möller AT, Kotzé HF, Sieberhagen KJ. Department of Psychology, University of Stellenbosch, RSA.
Jump up ^ Pratkanis, A. R. (1992). "The cargo-cult science of subliminal persuasion". Skeptical Inquirer. 16: 260–272.
Jump up ^ Greenwald, A.G.; E. R. Spangenberg; A. R. Pratkanis; J. Eskenazi (1991). "Doubleblind tests of subliminal self-help audiotapes". Psychological Science. 2: 119–122. doi:10.1111/j.1467-9280.1991.tb00112.x.
Jump up ^ Auday, B.C.; J. L. Mellett; P. M. Williams (1991). "Self-improvement Using Subliminal Selfhelp Audiotapes: Consumer Benefit or Consumer Fraud?". Psychological Science: Paper presented at the meeting of Western Psychological Association, San Francisco, CAlif., April.
Jump up ^ Moore, T.E. (1995). "Subliminal self-help auditory tapes: An empirical test of perceptual consequences". Canadian Journal of Behavioural Science. 27 (1): 9–20. doi:10.1037/008-400X.27.1.9.
^ Jump up to: a b Smith, Kirk H.; Rogers, Martha (1994). "Effectiveness of subliminal messages in television commercials: Two experiments". Journal of Applied Psychology. 79 (6): 866–874. doi:10.1037/0021-9010.79.6.866.
^ Jump up to: a b c Karremans, J.; Stroebe, W.; Claus, J. (2006). "Beyond Vicary's fantasies: the impact of subliminal priming and brand choice☆". Journal of Experimental Social Psychology. 42 (6): 792–798. doi:10.1016/j.jesp.2005.12.002.
Jump up ^ Cooper, Joel; Cooper, Grant (2002). "Subliminal motivation: A story revisited". Journal of Applied Social Psychology. 32 (11): 2213–2227. doi:10.1111/j.1559-1816.2002.tb01860.x.
Jump up ^ Key, W.B. (1973), Subliminal seduction: Ad media's manipulation of a not so innocent America, Englewood Cliffs, NJ: Prentice-Hall, ISBN 0-13-859090-7
Jump up ^ Dijksterhuis, Ap; Smith, Pamela K.; van Baaren, Rick B.; Wigboldus, Daniel H.J. (2005). "The unconscious consumer: Effects of environment on consumer behavior". Journal of Consumer Psychology. 15 (3): 193–202. doi:10.1207/s15327663jcp1503_3. CS1 maint: Uses authors parameter (link)
Jump up ^ Milliman, Ronald E. (1982). "Using background music to effect the behavior of supermarket shoppers". Journal of Marketing. 46 (3): 86–91. doi:10.2307/1251706.
Jump up ^ "Section 3. Misleading-advertising". BCAP Code. Committees on Advertising Practice (CAP). 2013. Retrieved 6 January 2014. "No advertisement may use images of very brief duration, or any other technique that is likely to influence consumers, without their being fully aware of what has been done."
Further reading[edit]
Boese, Alex (2006), Hippo Eats Dwarf: A Field Guide to Hoaxes and Other B.S., Orlando: Harcourt, pp. 193–195, ISBN 0-15-603083-7
Dixon, Norman F. (1971), Subliminal Perception: The nature of a controversy, New York: McGraw-Hill, ISBN 0-07-094147-5
Greenwald, A. G. (1992). "New Look 3: Unconscious cognition reclaimed". American Psychologist. 47 (6): 766–779. PMID 1616174. doi:10.1037/0003-066X.47.6.766.
Holender, D. (1986), "Semantic activation without conscious identification in dichotic listening, parafoveal vision, and visual masking: A survey and appraisal", Behavioral and Brain Sciences, 9 (1): 1–23, doi:10.1017/s0140525x00021269
Merikle, P. M.; Daneman, M. (1998), "Psychological Investigations of Unconscious Perception", Journal of Consciousness Studies, 5 (1): 5–18
United States Senate, Ninety-ninth Congress, First Session on Contents of Music and the Lyrics of Records (September 19, 1985), Record Labeling: Hearing before the Committee on Commerce, Science, and Transportation, Washington, DC: U.S. Government Printing Office CS1 maint: Multiple names: authors list (link)
Watanabe, T.; Náñez, J.; Sasaki, Y. (2001). "Perceptual learning without perception". Nature. 413 (6858): 844–848. PMID 11677607. doi:10.1038/35101601.
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