Cellular Senesence 42 min read January 28, 2026

What is Efferocytosis?

## When Dead Cells Don’t Get Cleaned Up, Diseases Emerge: “This process, termed efferocytosis, is critical for the prevention of autoimmune and inflammatory disorders, and persistence of dead cells in tissue is characteristic of many human autoimmune diseases, notably systemic lupus erythematosus.” (3) “Aberrations in efferocytosis are associated with numerous inflammatory pathologies, including atherosclerosis, cancer and infections. The recent exciting discoveries defining the molecular ma

What is Efferocytosis?

by Dr. Timothy J.

Definition of Efferocytosis:

“In cell biology, _efferocytosis_ is the process by which apoptotic cells are removed by phagocytic cells. It can be regarded as the 'burying of dead cells'.” (1)

“Efferocytosis is a term derived from the Greek (meaning to carry the dead to the grave), and refers to the phagocytic engulfment of a cellular corpse. Both professional phagocytes (e.g. macrophages) and non-professional (e.g. neighboring) cells participate in efferocytosis.” (2)

Proper Efferocytosis is a Silent, Non-Inflammatory Process.

“Efferocytosis is a carefully orchestrated process by which phagocytes are recruited to sites of cell death, recognize and engulf dying cells, and clear them in an 'immunologically silent' manner.”

“The most notable characteristic of the efferocytosis of apoptotic cells is its ‘immunologically silent' response. Although the mechanisms by which phagocytes facilitate engulfment of dead cells has been a well-studied area, the pathways that coordinate to process the ingested corpse and direct the subsequent immune response is an area of growing interest. The recently described pathway of LC3 (microtubule-associated protein 1A/1B-light chain 3)-associated phagocytosis (LAP) has shed some light on this issue. LAP is triggered when an extracellular particle, such as a dead cell, engages an extracellular receptor during phagocytosis, induces the translocation of autophagy machinery, and ultimately LC3 to the cargo-containing phagosome, termed the LAPosome. In this review, we will examine efferocytosis and the impact of LAP on efferocytosis, allowing us to reimagine the impact of the autophagy machinery on innate host defense mechanisms.” (3)

When Dead Cells Don’t Get Cleaned Up, Diseases Emerge:

“This process, termed efferocytosis, is critical for the prevention of autoimmune and inflammatory disorders, and persistence of dead cells in tissue is characteristic of many human autoimmune diseases, notably systemic lupus erythematosus.” (3)

“Aberrations in efferocytosis are associated with numerous inflammatory pathologies, including atherosclerosis, cancer and infections. The recent exciting discoveries defining the molecular machinery involved in efferocytosis have opened many avenues for therapeutic intervention, with several agents now in clinical trials.” (5)

History of Apoptosis

“Central to immune cell function is the removal of apoptotic cells, termed “efferocytosis”. In non-resolving diseases, particularly atherosclerosis, efferocytosis declines, areas of necrosis expand, and robust inflammatory responses arise. Despite the multiple methods that lower circulating levels of cholesterol, atherosclerotic cardiovascular disease remains the leading cause of death worldwide. The recent success of the CANTOS and COLCOT clinical trials indicate that lowering inflammation is a critical arm of the treatment strategy to decrease the burden of atherosclerosis. However, anti-inflammatory therapies leave individuals at risk for acute infections due to a reduced ability to mount an immune response. Because efferocytosis instead activates pro-resolving pathways while leaving acute inflammatory responses intact, enhancing the clearance of dead cells becomes an attractive therapeutic strategy to curb atherosclerosis. However, several areas related to efferocytosis remain unsolved. These include how macrophages handle the overwhelming burden of nutrients after internalizing an AC, what are the processes continuing the successive clearance of apoptotic cells, and what pathways are in place that transition the termination of inflammation into resolution. These topics are the focus of this Review. A timeline of significant events in the efferocytosis field are shown.” (7)

Efferocytosis Overview: Find Me, Eat Me, Post-Meal

"Clearance processes for apoptotic cell by phagocytes rely on “find me” and “eat me” signals that signal their internalization:

• (A) “Find me” signal: apoptotic cells release signals that attract phagocytes to the site of programmed cell death. These signals include nucleotides (UTP and ATP), S1P, CX3CL1, and LPC. Phagocytes recognize the “find me” signals using cognate receptors such as P2Y2, S1PRs, CXCR3, and G2A.

• (B) “Eat me” signal: the apoptotic cells express “eat me” signals that allow phagocytes to recruit surface receptors and bridging molecules to identify and engulf apoptotic cells. PtdSer is a primary “eat me” signal expressed by phagocytes and is directly recognized by phagocytic receptors including BAI, TIM4, and stabilin 2. Phagocytes can employ avB3 and tyrosine kinase receptor such as Mertk bind to PtdSer indirectly through bridging molecules such as MFG-E8 and Gas-6, respectively.

• (C) The engulfment process: after recruitment of engulfment receptors via the CRKII-ELMO-DOCK180 complex within phagocytes, rac-1 signaling pathway is activated for phagosome formation. Once the phagosome is formed, LC3 is recruited to phagosomes (now the LAPosome) leading to lysosome-mediated digestion of the internalized apoptotic body. Degraded products release fatty acids that stimulate LXR and PPARγ for cholesterol efflux leading to the production of anti-inflammatory cytokines such as TGF-β, IL-10 and IL-13.” (8)"

Efferocytosis In Depth, The 4 Stages:

"Macrophages bind externalized phosphatidylserine (PtdSer) on apoptotic cells using cell surface receptors, such as the TIM family of immunoglobulin receptors, Stabilin 1 or 2, or the GPCR BAI1. Alternatively, macrophages bind apoptotic cells indirectly through bridging molecules, such as Gas6 or proteins. Activation of cell-surface receptors stimulate Rac1 activation at the site of apoptotic cell adhesion to drive formation of the phagosome and mediate internalization. LC3-associated phagocytosis occurs through a Rubicon-dependent manner that acilitates lysosome fusion to promote apoptotic cell degradation. Completion of efferocytosis stimulates the production of anti-inflammatory cytokines and synthesis of SPMs to drive nflammation resolution.” (7)

LC3-LAP is the Bridge Between Efferocytosis and Autophagy

“Efferocytosis is a term coined by Peter Henson and colleagues in 2003 to describe the engulfment and clearance of cells undergoing apoptosis by phagocytes. Defects in efferocytosis can result in the rupture of uncleared apoptotic bodies and the subsequent leakage of cytosolic contents onto adjacent cells and exposure of tissues to oxidants, harmful enzymes and factors including caspases and other proteases. This can induce inflammation and tissue damage through a process referred to as secondary necrosis. Less understood is that during efferocytosis, signals from neighboring cells can initiate particular components of canonical autophagy to engage with the phagosome to increase the efficacy of the digestion of engulfed cellular cargo.

When components of autophagy are recruited to cargo-containing single membrane phagosomes to form LAPosome for digestion and clearance, the process is called LAP. Hence, LAP is a form of non-canonical autophagy that represents a hybrid between the processes governing the autophagic clearance of intracellular factors and those that accept extracellular cargos for degradation.”

“As the name suggests, a distinguishing feature of LAP is the recruitment of microtubule-associated protein light chain 3 (LC3) to the phagosome membrane after internalization of apoptotic cells or pathogens. In the normal situation, LAP is vital to resolve inflammation, promote wound healing and prevent auto-immunity and immune-mediated tissue damages. Reports by Martinez et al. show that dysregulation of LAP impairs efferocytosis, culminating in necrosis of the phagocyte that has engulfed the uncleared cell and uncontrolled inflammation. These findings suggest that LAP may be essential for efferocytosis and preventing the many deleterious consequences of secondary necrosis. In support of this, Cunha and colleagues demonstrated that defects in LAP restricts tumor immunosuppression afforded by efferocytosis. Hence, these studies provide evidence that LAP may also be an influential contributor to the production of tolerogenic signals that are activated upon engulfment of apoptotic cells during efferocytosis. Martinez et al. further showed that in the absence of LAP, phagocytosis of apoptotic bodies still occurs but the vesicles remain in the phagocyte for a prolonged duration or are not digested, and points to a critical role of LAP in finalizing the intracellular digestion process. Hence, in the context of cancer, defects in LAP have paradoxically beneficial effects including the activation of tumor infiltrating lymphocytes (TIL), enhanced effector T cells activity and elevated secretion of inflammatory mediators including stimulator of interferon genes (STING), type I interferons (IFNs) and tumor necrosis factor alpha (TNF-α). Taken together, these data suggest that, (1) LAP is critical for governing the digestion of the engulfed cellular debris and prevention of inflammatory and immune responses, (2) internalization of dead or dying cells alone is not solely responsible for immunosuppressive activities in the context of clearing apoptotic cells, and (3) LAP activation plays a critical role in immune tolerance. These recent findings of LAP as a concomitant effector of efferocytosis brings new significance to the processes that govern the clearance of apoptotic cells in the tumor microenvironment (TME), where there is frequent instances of programmed cell death”

“LC3-Associated phagocytosis (LAP). LAP recruitment process is triggered for degradation of the phagocytosed cargo following phagosome formation. The cargo is recognized by cell recognition receptors such as TIM-4 which causes the cargo to be engulfed in a single membrane phagosome called LAPosome. This process is initiated by recruitment of PI3KC3 complex consisting of Rubicon, vps34, beclin-1, and vps15 which enable PI3P to be localized to the LAPosomes. This stabilizes the NOX2 complex to produce ROS which is necessary to recruit LC3-II to the phagosome (LAPosome). The LAPosome then fuses with lysosomes to mature and to effective e digest the cargo.” (8)"

Senescent Cells Suppress Efferocytosis

“Senescent cell-mediated efferocytosis suppression (SCES) was independent of the senescence-associated secretory phenotype (SASP) but instead required direct contact between macrophages and senescent cells.” (4)

“The daily removal of billions of apoptotic cells in the human body via the process of efferocytosis is essential for homeostasis. To allow for this continuous efferocytosis, rapid phenotypic changes occur in the phagocytes enabling them to engulf and digest the apoptotic cargo. In addition, efferocytosis is actively anti-inflammatory and promotes resolution. Owing to its ubiquitous nature and the sheer volume of cell turnover, efferocytosis is a point of vulnerability.” (5)

“Efferocytosis is the process of rapid and efficient clearance of apoptotic cells by both professional and non-professional phagocytic cells.”

“Removal of apoptotic cells through efferocytosis prevents secondary necrosis and the resultant inflammation caused by the release of intracellular contents. The importance of efferocytosis in homeostasis is underscored by the large number of inflammatory and autoimmune disorders, including atherosclerosis and systemic lupus erythematosus, that are characterized by defective apoptotic cell clearance. Although mechanistically similar to the phagocytic clearance of pathogens, efferocytosis differs from phagocytosis in that it is immunologically silent and induces a tissue repair response. Efferocytes face unique challenges resulting from the internalization of apoptotic cells, including degradation of the apoptotic cell, dealing with the extra metabolic load imposed by the processing of apoptotic cell contents, and the coordination of an anti-inflammatory, pro-tissue repair response.” (6)

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Posted in: Cellular Senesence

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