13- Effector responses: antibody and cell mediated immunity
Once B cells and T cells are activated, how do they kill pathogens and cancer cells?
They constantly patrol and target an immune response.
Learning Objectives
Explain how humoral immune responses protect us against pathogens, toxins, or malignancy.
For antibody responses, recognize how the multiple immunoglobulin classes and subclasses can have common, but also distinct, mechanisms for antigen inactivation, elimination, tissue distribution, and other biological activities.
Describe the multistep process required for inducing the differentiation of TC precursors into effector cytotoxic T lymphocytes (CTLs).
Compare and contrast the mechanisms by which CTLs and natural killer (NK) cells recognize and kill infected or tumor target cells.
Explain how NK cells and NKT cells each have some properties of innate immune cells and adaptive immune cells.
Key Terms
Humoral Immunity: antibodies target extracellular pathogens using class dependent effector functions
Cell-mediated Immunity: cells target intracellular and extracellular pathogens as well as abnormal cells using apoptosis and degranulation
Innate cells: NKT, NT, neutrophils, macrophages, eosinophiles
FcR: Receptors on the surface of cells that recognize the constant fragments of antibodies
neutralization: inactivates pathogens and prevents it binding to target cell
agglutination: crosslinks multiple pathogens because antibodies are multivalent
opsonization: marks pathogen for macrophage phagocytosis; macrophage contains Fc receptor that will bind to constant fragment of antibody
complement cascade activation: lysis of bacteria if MAC becomes activated or opsonization
Antibody cell-mediated cytotoxicity: NK cells bind to peptides and phagocytose
degranulation and mediator release: granulocytes (esp. eosinophiles and mast cells) binds to antibody, releases granules and kills pathogen
Natural antibodies: B1B T cell independent antigen nonconventional cells
Transcytosis: transcellular transport of antibodies across interior of a cell captured by vesicles on one side and ejected on other side
ITAM: immunoreceptor tyrosine activation motif; positive signaling
ITIM: immunoreceptor tyrosine inhibition motif; negative signaling
Myeloid lineage: granulocytes and mono, macro and baso/mast and dendritic cells (innate- BM maturation)
Lymphoid lineage: lymphocytes (B and T) dendritic cells, and ILC (NK, NKT) (adaptive-lymphoid organ maturation)
PolyIgR: polymeric immunoglobulin receptor is expressed by epithelial cells that initiate transcytosis of IgA and IgM from blood to lumen of tissues
Neonatal FcR: transports IgG into fetus, from placenta, blood and into mucosal secretions
Monoclonal Antibodies mAb: mouse derived antibodies
Chimeric antibodies: mouse variable regions, human constant regions
Humanized antibodies: CDRs are mouse derived
Human antibodies: entirely human regions
Rituximab: binds to CD20 (B-cell specific antigen); destroys B cells through complement MAC or CDCC activation
Perforin: protein that creates pores in the cell membrane to allow granzyme inside
Granzyme: serine protease induces apoptosis by activating Caspase signaling pathway
Fas-FasL Pathway: This pathway involves the interaction between Fas, a death receptor on target cells, and Fas ligand (FasL) expressed on activated T cells, leading to apoptosis of the target cell.
FADD: Fast activating Death domain that triggers the downstream signaling cascade necessary for apoptosis when Fas is engaged.
Notes
Effector responses target pathogens and abnormal cells through two immunity responses:
Humoral (antibodies)
EXTRACELLULAR PATHOGENS
CLASS DEPENDENT ON Ig GAMED
Function: isotype dependent
Cell-mediated (Th and CD8+)
INTRACELLULAR , EXTRACELLULAR AND ABNORMAL CELLS
Helper T cells, CD8, NK, NKT, and Innate cells (Neutrophils, macrophages, dendritic cells, and eosinophils and monocytes)
Function: apoptosis or degranulation
FCR (Constant fragment (region) of antibodies- receptors) link both immunity responses by optimizing all cells that act as markers for phagocytosis
Effector functions of antibodies
neutralization: inactivates pathogens and prevents it binding to target cell
agglutination: crosslinks multiple pathogens because antibodies are multivalent
opsonization: marks pathogen for macrophage phagocytosis; macrophage contains Fc receptor that will bind to constant fragment of antibody
complement cascade activation: lysis of bacteria if MAC becomes activated or opsonization
Antibody cell-mediated cytotoxicity: NK cells bind to peptides and phagocytose
degranulation and mediator release: granulocytes (esp. eosinophiles and mast cells) binds to antibody, releases granules and kills pathogen
Each class of immunoglobulin will tailor specific pathogens.
Immunoglobulin | Function | Notes |
IgM | complement activation and agglutination (opsonization and phagocytosis) | pentameric molecule; low affinity antibodies (naive cells), high avidity (compensates for low affinity) |
IgG | neutralization and complement activation, opsonization and ADCC | most common; multiple subclasses 1-4 |
IgA | NOT pro-inflammatory (no complement activation); neutralization in secretion and opsonization in serum | dimer joined by J chain; two subclasses in serum and secretions, important in MALT (skin and mucosa surfaces) , help tolerate nonpathogenic bacteria good in gut |
IgE | degranulation of eosinophils; releases histamine | allergy and asthma; parasitic multicellular pathogens; low in serum; |
IdD | degranulation | terrible at everything; low in serum; in upper respiratory tract |
FcR Receptors- Antibody sensors
recognize constant fragments of antibodies
mediate effector functions of antibodies
link humoral and cell-mediated immune responses
allows nonspecific (innate) immune cells to take advantage of antigen specific (adaptive) antibodies
immunoglobulin family domain
Majority: Positive ITAMS
Minority: Negative ITIMS (Fc gamma RIIB)
Differentiate by
Antibody class binding
Cell expression
Responses triggered
Multiple FcRs must be cross linked to initiate signal
Most diverse is Fc gamma because IgG IS MOST DIVERSE (1-4 SUBCLASSES)
Fcgamma : IgG (1-4)
Fcalpha : IgA
Fcmu: IgM
Fcepsilon: IgE (high and low)
Fcdelta: IgD
PolyIgR: polymeric immunoglobulin receptor is expressed by epithelial cells that initiate transcytosis of IgA and IgM from blood to lumen of tissues
carries Ab into tears and milk and gut mucosa with IgA and IgM
FcRn: neonatal Fc Receptor
transports IgG from placenta into fetus, milk in intestine into blood and into mucosal secretions
epithelial and endothelial cells
Making Connections
B cells encounter Ag, bind and become activated
Two types of B cells:
T independent : B1B and Marginal zone; TI-1 and TI-2
T dependent: B2B; Need Helper T cells to differentiate into different isotypes based on cytokines
Differentiate into:
Phase 1:
GC- independent memory B cell: low affinity
Short Lived plasma cell (sLPC): low affinity
GC- dependent memory B cell
Phase 2:
GC dependent memory B cells undergo SHM and class switching to become higher affinity
GC-dependent memory cells
Long lived plasma cells lLPC
AID: enzyme induces SHM
Dark to light for affinity selection testing on fDC
centroblasts → centrocytes
B cells become Ag secreting plasma cells
intracellular pathogens: class switch to isotypes that activate ADCC
extracellular pathogens: class switch to IgG subclasses
AB is carried to mult. sites for isotype specific effector functions that is sensed by FcRs on target cells
Therapeutic uses of Antibodies
We can manufacture antibodies with the specific protein/antigen and inject the antigen into a lab animal and wait for the organism to develop antibodies for that specific protein/antigen and harvest their B cells to culture in the lab.
However, we can’t inject ourselves with mouse/pig/horse/animal antibodies because our bodies will detect them as foreign OR detect ourselves as foreign and develop an immune response against it. SO, we can engineer modifications to make them look like human antibodies.
Monoclonal antibodies mAb: antibodies of mouse origin make up all therapeutic antibodies that end in -mab.
Three types:
Chimeric: V (mouse) C (Human)
Humanized: CDRs (mouse)
Human: entirely human
Therapies:
Antagonism- blocks ligand or receptor from binding; no activation
Agonistic: antibody mimics ligand or receptor to activate pathway
Complement activation: promotes lysis of target cell by activating the complement
ADCC: antibody binds cancerous cells and T cells kill that cell
Examples:
Rituximab: binds to CD20 (B-cell specific antigen); destroys B cells through complement MAC or CDCC activation (marks them for destruction)
B cell cancer; Non-Hodgkin’s lymphoma
B cells replenished by bone marrow and hematopoietic stem cells
Cell Mediated Effector Responses
Cytotoxic effector cells: Adaptive, Branch and Innate Immunity arms
Apoptosis kills infected cells and abnormal cells
CTLs and NK involved in organ rejection
CD8+
CD8+ recognize infected and tumor cells by TCR activation on the cell surface of the T cell by binding MHC I peptide complex on target cell, local IL-2 cytokine signaling and costimulatory signals from CD28-CD80/86 interaction between pAPC and T cell
The CD8+ cells are activated by pAPC that have been licensed by CD4+ cells for crosspresentation
How do these cells kill target cells? They secrete cytotoxic granules containing perforin and granzyme inducing Caspase signaling pathway and Fas-FasL pathway
Fas-FasL pathway: Fas ligand on CD8+ cell membrane binds to Fas receptor on target cell, activates fast activating death domain (FADD) and cleaves Procaspase into caspase, and active Caspase 8 will trigger apoptosis of target cells.
NK- Natural Killer Cells
Innate lymphoid cell
Lack Ag specific receptors- are the first wave of immunity before CD8+ adaptive cells to respond to infected cells
Activated by IFN-alpha, beta and IL-2
Recognize inhibitory and activating receptors on target cells through two models: Missing MHCI self and Balance signal model
Viruses decrease expression of MHC I to try to evade CD8+ Tc cells, however, NK cells recognize the lack of self
Must be licensed and tested
NK cell memory exists despite it being innate cell
NKT- Natural Killer T cell
Bridge
Share characteristics of both
conserved TCR
Fas-FasL
no memory
Questions
Match each antibody function to the correct definition:
Neutralization: most effective mode of protection from toxins and viruses
Agglutination: binds and crosslinks multiple antigens
Opsonization: enhances the engulfment of antigens by phagocytes
Complement activation: results in formation of the MAC cascade
ADCC: activates the killing activity of cytotoxic cells including NK cells
Match each description with correct antibody isotype:
IgA: found in secretions; effective at neutralizing toxins and pathogens
IgM: first antibody produced in primary response'; pentavalent
IgG: includes several subclasses, some of which are particularly good at complement fixation
IgD: promotes basophil and mast cell release of AMPs
IgE: best known for roles in allergy, asthma, and parasitic worm infection
What are the two types of immunity? Humoral and cell mediated
Which immunity type targets extracellular pathogens only? humoral
Which immunity type targets pathogens AND abnormal cells? cell-mediated
What effector functions does cell-mediated immunity perform? degranulation and apoptosis
What effector functions does humoral immunity perform? class-dependent because they’re antibodies Ig GAMED
Cell-mediated immunity is mediated with the help of which adaptive cells? Helper T cells (CD4+) and CD8+ cytotoxic t cells
Cell-mediated immunity is mediated by which innate cells? NKT, NK, and neutrophils, macrophages, and eosinophiles
How does cell-mediated and humoral immunity interact with each other? FcRs: Constant fragment (region) of antibodies- receptors
How do FcRs work? How does it link both immunity responses? Antibodies will opsonize a pathogen and the FcRs on the innate/adaptive cells will recognize the constant region of the antibodies and that will facilitate the phagocytosis/degranulation of the pathogen
How many effector functions of antibody mediated immune responses are there? What are they? 6; neutralization, agglutination, opsonization, complement activation ADCC, degranulation
Match the antibody with its effector function (from the 6):
IgG: neutralization, MAC, opsonization, ADCC
IgA: neutralization, opsonization, ADCC, degranulation
IgM: agglutination, MAC
IgE: degranulation
IgD: degranulation
What is the relationship between avidity and affinity? inverse; compensates for low affinity
How does class switching in antibodies work to change isotypes? depends on the cytokines released from helper T cells
What type of B cells bear IgM isotype? B1B T cell independent antigen cells (nonconventional)
when are IgM antibodies released in B cell activation? Phase 1; germinal center independent
Which antibodies are produced first? IgM
The majority of IgM in circulation are called? natural antibodies; B1B cells
Which functions are performed by IgM antibodies? complement (MAC) activation and agglutination
Where are isotypes determined on the antibody structure? Heavy chain constant region
Which isotype is “terrible”? IgD
Which isotype is confined to the upper respiratory tract? IgD
Which isotype is pentameric? IgM
Which isotype is responsible for tolerating nonpathogenic bacteria in the gut? IgA
Which isotype is a dimer joined by J chain? IgA
Which isotypes have subclasses? IgA (1 and 2) & IgG (1-4)
Which isotypes have only 1 effector function? Which function is it? IgE and IgD; degranulation
Which isotype is found primarily on mucosal surfaces of barriers? IgA
Which isotype is known for role in allergies/asthma? IgE
Which isotype targets parasitic worms? IgE
Which isotype is most common in serum? IgG
Which isotype has the longest half-life? IgG
Which isotype is the heaviest? IgM (pentameric) and IgA (dimer)
Which isotype is the best at complement MAC activation? IgG
Which isotype crosses placenta? IgG
Which isotype is secreted? IgA
Which cells secrete IgA? Plasma cells
What are the Fc Receptors on epithelial cells for IgA called? poly-Ig receptor
What is transport of an antibody from one domain into another called? Transcytosis
To which superfamily do Fc receptors belong to? Immunoglobulin domain family
The majority of FcR are part of which sub-family? R
The majority of FcR are (inhibitory/activating)? activating
Which FcR is the only inhibitory receptor? Why? Fc gamma RIIB- has ITIM
Which FcR is the most diverse? Fc gamma R
Which cells activate phagocytosis? macrophages
Which cells induce degranulation? granulocytes (Neutrophils, Eosinophils, Basophils, Mast cells)
Which cells activate ADCC (antibody dependent cell-mediated cytotoxicity)? NK cells
Fc gamma RIIB is an inhibitory FcR. How does it inhibit? blocks B cell activation and proliferation
When is Fc gamma RIIB expressed? when there are enough antibodies in circulation as a negative feedback regulatory mechanism
Which Fc epsilon R is high affinity? FceRI (one)
Which Fc epsilon R is low affinity? FceRII (two)
Which Fc epsilon R has unknown function? FceRII
Fc epsilon R triggers what effector function? Histamine, protease, inflammatory molecule release of IgE cells
Fc mu R are expressed by which cells and perform which function? B/T and NK cells; regulate cell activation
Fca/Fcm receptors are expressed on which cells? What function do they mediate? B, macrophages and fDC; binds both with high and low affinity to induce phagocytosis in IgM
Which FcR is responsible for carrying antibodies from the blood into the lumen of tissues? Polygenic Immunoglobulin Receptor
Why is poly-Ig receptor important? It carries antibodies from blood into lumen of tissues for tear, milk and gut mucosa secretion of IgA and IgM
Which isotypes are transcytosed through mediation of poly-Ig receptor? IgM and IgA
Which FcR is responsible for transporting IgG into the fetus? FcRn neonatal receptor
What are the antibodies for therapeutical uses called? Monoclonal antibodies mAb
What are the three types of mAB? chimeric, humanized and human
What are the 4 types of therapies using antibodies? antagonism, agonism, complement MAC activation, and ADCC
What does Rituximab target? B cell antigen CD20
cell mediated effector responses include which 3 “arms”? adaptive, innate and bridge
Which cells exhibit properties of both adaptive and innate responses in cell-mediated immunity? NKT
Which adaptive cell carries out cell mediated effector responses more commonly? CD8+ Cytotoxic T cells
Which cells are involved in organ/graft rejection? CTLs (CD8+) and NK
How do CD8+ respond to organ rejection? tissue not matched to the host MHC is killed by CD8+ cytotoxic granules containing perforin and granzyme
How do CD8+ cells recognize target cells? TCR activation (CD8 is a cytotoxic cell) by MHC1-peptide complex
Which MHC does a CD8+ recognize? MHC I presenting self antigens
How are dendrites licenced? Why do they need to be licensed? Licensed by engaging with CD4+ Th and to gain ability for cross presentation
What are the two models of cross presentation? sequential and simultaneous
What are the two pathways that CD8+ instruct apoptosis? Perforin/Granzyme AND Fas-FasL pathways
Why do we have CD8+ and NK cells if they perform similarly? What are the smiliarites and differences? Both kill infected cells; CD8+ are part of the adaptive immunity and NK are part of innate immunity
Which interferons are related to viral infections? IFN- alpha and Beta
NKs proliferate in response to? IFNa, IFN-B, and IL-12
How do NK cells recognize target cells? Missing self MHCI or balance signal models
Which cells must be licensed to perform functions? DC and NK
True/False: NK cell memory exists True, even though it’s innate cell, there is some memory