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Endocytosis and endosomal trafficking of plasma membrane proteins for degradation regulate cellular homeostasis and development. As part of these processes, ubiquitinated plasma membrane proteins (cargo) are recognized, clustered, and sorted into intralumenal vesicles of multivesicular endosomes by ESCRT (Endosomal Sorting Complexes Required for Transport) proteins. At endosomes, ESCRT proteins recognize ubiquitinated cargo and mediate the deformation of the endosomal membrane in a negative geometry, away from the cytosol. ESCRTs are organized in five major complexes that are sequentially recruited to the endosomal membrane where they mediate its vesiculation and cargo sequestration. ESCRTs also participate in other membrane remodeling events and are widely conserved across organisms, both eukaryotes and prokaryotes. Plants contain both conserved and unique ESCRT components and show a general trend toward gene family expansion. Plant endosomes show a wide range of membrane budding patterns with potential implications in cargo sequestration efficiency, plant development, and hormone signaling. Understanding the diversification and specialization of plant ESCRT proteins can provide valuable insights in the mechanisms of ESCRT-mediated membrane bending. In this review, we discuss the endosomal function of ESCRT proteins, their unique features in plants, and the potential connections to the modes of plant endosomal vesiculation. 

Multivesicular endosomes (MVEs) sequester membrane proteins destined for degradation within intralumenal vesicles (ILVs), a process mediated by the membrane-remodeling action of Endosomal Sorting Complex Required for Transport (ESCRT) proteins. InArabidopsis, endosomal membrane constriction and scission are uncoupled, resulting in the formation of extensive concatenated ILV networks and enhancing cargo sequestration efficiency. Here, we used a combination of electron tomography, computer simulations, and mathematical modeling to address the questions of when concatenated ILV networks evolved in plants and what drives their formation. Through morphometric analyses of tomographic reconstructions of endosomes across yeast, algae, and various land plants, we have found that ILV concatenation is widespread within plant species, but only prevalent in seed plants, especially in flowering plants. Multiple budding sites that require the formation of pores in the limiting membrane were only identified in hornworts and seed plants, suggesting that this mechanism has evolved independently in both plant lineages. To identify the conditions under which these multiple budding sites can arise, we used particle-based molecular dynamics simulations and found that changes in ESCRT filament properties, such as filament curvature and membrane binding energy, can generate the membrane shapes observed in multiple budding sites. To understand the relationship between membrane budding activity and ILV network topology, we performed computational simulations and identified a set of membrane remodeling parameters that can recapitulate our tomographic datasets. 

Endocytosis, secretion, and endosomal trafficking are key cellular processes that control the composition of the plasma membrane. Through the coordination of these trafficking pathways, cells can adjust the composition, localization, and turnover of proteins and lipids in response to developmental or environmental cues. Upon being incorporated into vesicles and internalized through endocytosis, plant plasma membrane proteins are delivered to the trans‐Golgi network (TGN). At the TGN, plasma membrane proteins are recycled back to the plasma membrane or transferred to multivesicular endosomes (MVEs), where they are further sorted into intralumenal vesicles for degradation in the vacuole. Both types of plant endosomes, TGN and MVEs, act as sorting organelles for multiple endocytic, recycling, and secretory pathways. Molecular assemblies such as retromer, ESCRT (endosomal sorting complex required for transport) machinery, small GTPases, adaptor proteins, and SNAREs associate with specific domains of endosomal membranes to mediate different sorting and membrane‐budding events. In this review, we discuss the mechanisms underlying the recognition and sorting of proteins at endosomes, membrane remodeling and budding, and their implications for cellular trafficking and physiological responses in plants.