Borregaard N, Lollike K, Kjeldsen L, Sengeløv H, Bastholm L, Nielsen M H, Bainton D F
Department of Hematology, University of Copenhagen, Rigshospitalet, Denmark.
Eur J Haematol. 1993 Oct;51(4):187-98. doi: 10.1111/j.1600-0609.1993.tb00629.x.
The traditional classification of neutrophil granules as peroxidase-positive (azurophil, or primary) and peroxidase-negative (specific or secondary) has proven to be too simple to explain the differential exocytosis of granule proteins and incorporation of granule membrane into the plasma membrane which is an important aspect of neutrophil activation. Combined subcellular fractionation and immunoelectron microscopy has revealed heterogeneity among both peroxidase-positive and peroxidase-negative granules with regard to their content, mobilization and time of formation. Peroxidase-negative granules may be classified according to their content of lactoferrin and gelatinase: 15% of peroxidase-negative granules contain lactoferrin, but no gelatinase. 60% contain both lactoferrin and gelatinase. The term specific or secondary granule should be reserved for these two subsets. In addition, 25% of peroxidase-negative granules contain gelatinase but no lactoferrin. These should be termed gelatinase granules or tertiary granules. Gelatinase granules are formed later than specific granules and mobilized more readily. In addition, a distinct, highly mobilizable intracellular compartment, the secretory vesicle, has now been recognized as an important store of surface membrane-bound receptors. This compartment is formed in band cells and segmented cells by endocytosis. This heterogeneity among the neutrophil granules is of functional significance, and may also be reflected in the dysmaturation which is an important feature of myeloproliferative and myelodysplastic disorders.
传统上把中性粒细胞颗粒分为过氧化物酶阳性（嗜天青颗粒或初级颗粒）和过氧化物酶阴性（特异性颗粒或次级颗粒），事实证明这种分类过于简单，无法解释颗粒蛋白的差异性胞吐作用以及颗粒膜并入质膜的过程，而这是中性粒细胞激活的一个重要方面。联合亚细胞分级分离和免疫电子显微镜技术已揭示，过氧化物酶阳性颗粒和过氧化物酶阴性颗粒在其内容物、动员和形成时间方面均存在异质性。过氧化物酶阴性颗粒可根据其乳铁蛋白和明胶酶的含量进行分类：15%的过氧化物酶阴性颗粒含有乳铁蛋白，但不含明胶酶。60%的颗粒同时含有乳铁蛋白和明胶酶。术语特异性颗粒或次级颗粒应保留用于这两个亚群。此外，25%的过氧化物酶阴性颗粒含有明胶酶但不含乳铁蛋白。这些应称为明胶酶颗粒或三级颗粒。明胶酶颗粒比特异性颗粒形成得晚，且更容易被动员。此外，一个独特的、高度可动员的细胞内区室，即分泌小泡，现已被确认为表面膜结合受体的重要储存部位。这个区室在杆状核细胞和分叶核细胞中通过内吞作用形成。中性粒细胞颗粒之间的这种异质性具有功能意义，也可能反映在发育异常上，而发育异常是骨髓增殖性疾病和骨髓增生异常综合征的一个重要特征。
