Lecture 22: Virus offence meets host defense: Innate Immunity
BSCI437
Flint et al. Chapter 15, pp. 531 – 584.
General points
• We literally swim in viruses and other microbes
• We’ve evolved numerous, overlapping active and passive defenses to enable us to
ward off infection
• Successful pathogens have evolved gene products that modify, redirect and block
host defenses
For every host defense, there is a viral offense.
Primary physical and chemical defenses
Skin
o Largest organ in body – >5 kg for adults
o Strong barrier to infection
o Inactivates viruses by desiccation, acids, skin cleansing mechanisms,
inhibitors made by commensals
Other Epithelial surfaces: inactivates viruses via
o Secretions
o Mucus
o Tears
o Acid pH
Intrinsic cellular defenses – Intro
Highly conserved, arose early in evolution
Non-specific responses to stresses, e.g.
Starvation
Irradiation
Infection
Apoptosis (Fig. 15.1)
Programmed cell death
Keeps “wrong” cells from proliferating
o “Sacrifice the one for the good of the many”
Promoted by Capsases: Cysteine proteases that cleave after ASPartate residues
o Extrinsic pathway: triggered from outside
o Intrinsic pathway: triggered from inside
Viruses can alter gene expression inside and outside of cell
Viruses can activate either pathway
Viral inhibition of apoptosis (see Fig. 15.2)
Many viral proteins have evolved to inhibit apoptosis by intervening at any one of
multiple steps in the pathway.
Other intrinsic defenses
Autophagy
o Cells cannibalize themselves!
o Induced by starvation, viral infection
Nuclear domain 10 (NSD10) bodies
o Prevent transcription of “foreign DNA” in nucleus
o Defense against DNA viruses
RNA silencing (siRNAs, RNAi)
o System recognizes and degrades dsRNAs
o Defense against RNA viruses
Immune defenses – overview (Fig. 15.3.)
Highly coordinated
Depends on interplay of secreted proteins, receptor-mediated signaling, and cellto-cell communication
Three critical steps: recognition, amplification, control.
Can be non-adaptive (innate) or adaptive.
Innate immune responses
Cytokines – soluble proteins that act as signals
Sentinel cells – e.g. dendritic cells, patrol local areas
Complement – soluble proteins, poke holes in infected cells
Cytolytic “death star” cells
o Natural Killer (NK) cells
o Neutrophils, Basophils, Macrophages, other granulocytes.
Pattern recognition receptors
o Detection of “non-self”
Toll-like receptors.
Expressed inside and outside of cells
Very ancient
o Detection of “missing” or “altered self”
Used mainly by NK T-cells.
Also ancient: similar systems used by insects.
Cytokines
Rapid response team: one of the first indicators of infection
Infected cells make cytokines in response to multiple cues
e.g. uncoating of viruses, exposure to foreign DNA/viral proteins, stress
such as ER overloaded viral proteins or too much transcription activity.
Divided into three classes:
Proinflammatory – promote immune activation
o e.g. IL-1, Tnf, IL-6, IL-12
Anti-inflammatory – return system to basal activity
o e.g. IL-10, IL-4, Tgf-
Chemokines – Recruit immune cells early in immune response
o e.g. IL-8
Cytokines – viral responses (see Tables 15.6 and 15.7)
Cytokines interact with cells via cytokine receptors
Many viruses have evolved gene products to modulate the immune response
Virokines: Mimic host cytokines. (Table 15.6)
Viroreceptors: mimic host cytokine receptors. Serve as sinks for cytokines,
prevent cytokine activities. (Table 15.7)
Interferons – early warning (Table 15.8)
Produced by infected cells (and immature dendritic cells)
IFN-: produced by most nucleated cells in response to viral infection, dsRNA
IFN-: produced by most nucleated cells in response to viral infection, dsRNA
INF-: Produced by T-cells, NK cells in response to antigens, mitogens, IL-2, 12
Induce antiviral states
o Apoptosis of infected cells
o Apoptosis of nearby cells – firebreak to infection
o Block cell proliferation
o Enhances NK cell activity
o Alters MHC expression…and many more effects
Very non-specific and toxic…Not the “magic bullet”
o However, useful for treatment of many persistent viral infections, e.g.
hepatitis B and C
Antiviral IFN-induced proteins
Pkr (dsRNA activated protein kinase)
Activated by dsRNA.
Phosphorylates eIF2-, shutting down translation.
Many viral proteins evolved to inactivate or misdirect Pkr.
RNase L + 2’-5’ Oligo(A) synthetase
Work together to degrade RNAs.
Both antiviral and apoptotic
Mx proteins
In mice, specifically prevents “cap-snatching” by influenza
In humans ,can also prevent replications of VSV, Measles, parainfluenza and
others.
P200 proteins
Block cell proliferation
Inhibit rRNA transcription and ribosome biosynthesis
Nitric oxide synthase
Directs synthesis of NO in NK cells
Cytotoxic
Inhibits poxvirus and herpesvirus replication
PML
Part of the ND10 bodies, prevents transcription of foreign DNA
Ubiquitin-proteosome pathway components
Proteins tagged with ubiquitin are targeted to the proteosome for degradation
Viral gene products that counter IFN response (see Table 15.9)
Viruses have evolved many gene products to counter IFN. Types of modulation
include:
Inhibition of IFN synthesis
IFN receptor decoys
Inhibition of IFN signaling
Block functions of IFN-induced proteins
Non-specific cellular response (Figs. 15.9, 15.10)
Dendritic cells and macrophages
o Sentinel cells: patrol local area for bad guys
o Phagocytotic
o Present foreign antigen to T-cells
NK cells
o Recognize and kill virus infected cells
o Detect “altered” or “missing-self” molecules
Altered self: MHC I & II + non-self antigen
Missing self: Lack of MHC I or II, lack of activating receptors.
Complement (Fig. 15.11)
“Complemented by antibody”
3 pathways: classical, alternative, mannan-binding
Poke holes in infected cells, target cells for degradation
Short Interfering RNAs (siRNA)
No cellular dsRNAs, but RNA viruses produce these
Ancient mechanism against parasitic dsRNAs.
Two host cellular enzyme complexes
1. Dicer: recognizes and cleaves dsRNAs into small pieces (20 – 23 nt)
2. RISC complex: anneals small RNAs to target (viral) RNA, and then
cleaves the target.
Viral defenses against siRNA
Best known examples are in plants
• Coat protein of Turnip Crinkle Virus binds to and inactivates Dicer (see Lect. 7).
• Carnation Italian ringspot virus (CIRV) p19 protein.
– Specifically binds 21 nt dsRNA fragments.
– Acts as a molecular caliper to specifically select siRNAs based on the
length of the duplex region of the RNA.
– Sequesters siRNA away from RISC complex