5

Haraldsson B., Nystrom J., Deen W. M. Properties of the Glomerular Barrier and Mechanisms of Proteinuria Dept. Mol. Clin. Med./Nephrol., Inst. Med., Goteborg Univ., Sweden; Depart. Chem. Engineer., MIT, Cambridge, Massachusetts Physiol. Rev. 88: 451–487, 2008

6

I. Введение

1 2 3 4

7 8 9

Гломерулярный барьер обеспечивает фильтрацию воды, низко- и среднемолекулярных ве­ ществ и почти полностью ограничивают фильтрацию альбумина и других крупных белков. У человека образуется около 180 л/сут первичной мочи.

16

В обзоре рассмотрены функции гломерулярного барьера, механизмы протеинурии и нефроти­ ческих синдромов. Рассмотрены функции эндотелия, его гликокаликса и базальной мембра­ ны. Рассмотрены взаимодействия между барьером и белками плазмы (альбумин и др.), основные компоненты гломерулярного барьера, скорость клубочковой фильтрации, скорость кровотока и концентрации альбумина. Рассмотрены методы изучения проницаемости клубоч­ ков, физические принципы и теоретические модели, отражающие функционирование гломе­ рулярного барьера.

17

II. Компоненты гломерулярного барьера

18

A. Подоциты

10 11 12 13 14 15

19 20 21 22

Подоциты расположены снаружи от гломерулярных капилляров, на поверхности капсулы клубочка и обращены в сторону первичной мочи. Эти клетки имеют большое тело и длинные покрытые мембраной выросты цитоплазмы. Компоненты мембраны ограничивают проницае­ мость фильтра для макромолекул (молекулярная масса, заряд, конфигурация).

24

Щели мембраны подоцитов препятствуют фильтрации альбумина и миоглобина (116, 144, 254, 296, 319, 330, 342, 348).

25

B. Базальная мембрана

23

33

Базальная мембрана состоит из сети волокон коллагена IV типа (цепи α3, α4 и α5) (193, 271), ламинина (в основном ламинина 11; α5βγ1) (205), нидогена/энтактина, протеогликанов агри­ на и перлекана, а также гликопротеинами. Базальная мембрана гломерулярных капилляров намного толще (240—370 нм) (163) базальной мембраны других сосудов (40—80 нм) (288). Коллаген IV типа формирует основу базальной мембраны. Мутации цепей коллагена вызыва­ ют гломерулонефрит (14, 170, 195) или истончение гломерулярной мембраны (320). Базаль­ ная мембрана содержит протеогликанов, связанных с цепями гепарансульфата, что также обеспечивает селективность барьера (65, 73, 118, 145).

34

C. Эндотелий

26 27 28 29 30 31 32

35 36 37 38 39 40 41

Эндотелиоциты клубочка плоские, расположенные вокруг капиллярных петель (288, 315). В капиллярах клубочка клетки имеют фенестры, составляющие 20—50 % поверхности эндо­ телия (40, 73). Диаметр фенестр составляет 60 нм (316), а размер молекулы альбумина — 3,6 нм. Таким об­ разом, фенестры не ограничивают фильтрацию альбумина. Эндотелиоциты имеют заряд-се­ лективные свойства посредством отрицательно заряженных протеогликанов (260) или других селективных молекул (11, 52, 70, 134, 135, 212, 267, 268, 302). Стр. 1 из 22

42 43 44 45 46 47

D. Поверхностный эндотелиальный слой Эндотелиоциты поверхностного слоя участвует в свертывании крови (16, 174), ангиогенезе (39,96), реологии (62) и являются капиллярным барьером (129,134,135,301). Этот слой состо­ ит из двух компонентов: гликокаликса и большего эндотелиального слоя (44, 63, 178). По­ верхностный слой состоит из отрицательно заряженных гликопротеинов, гликозаминоглика­ нов, мембранных и секретируемых протеогликанов.

49

Отрицательно заряженные структуры эндотелия всего способствуют избирательной проница­ емости гломерулярной стенки (77, 112, 114, 115, 117, 121, 135, 278, 279, 289, 339).

50

1. Гликокаликс подоцитов

48

57

Подоциты покрыты гликокаликсом из сульфатированных гликозаминогликанов, сиализиро­ ванных гликоконьюгатов и гепарансульфата (232, 261). Основной сиалопротеин подоцитов подокаликсин сильно гликозилирован (151). У крыс, пуромицин снижал силовых кислот в подокаликсине и величину отрицательного заряда мембраны подоцитов (152). Гломеруляр­ ные клетки продуцируют протеогликаны глипикан-1 и синдикан-4 (25). Отрицательный заряд способствует сохранению расстояния между париетальными и висцеральными эпителиоци­ тами, что обеспечивает сохранение структуры и функцию гломерул (150).

58

E. Мезангий

51 52 53 54 55 56

70

Мезангий поддерживает структуру и функцию гломерулярного барьера. Мезангий состоит из мезангиальных клеток и внеклеточного матрикса. Мезангиальные клетки способны сокра­ щаться (162, 277). Сократимость может регулировать растяжимость гломерул в ответ на изме­ нение давления. Мезангиальные клетки продуцируют компоненты мезангиального матрикса, который состоит из коллагена (I, III, IV, V типов), ламинина, фибронектина и протеогликанов с цепями гепарансульфата и хондроитинсульфата (189, 215). Мезангиальные клетки выделя­ ют IL-1, фактор роста тромбоцитов (PDGF) и инсулиноподобный фактор роста (IGF) (1, 8. 352). PDGF стимулирует митоз клеток мезангия. Трансформирующий фактор роста (TGF)-β ингибирует клеточную пролиферацию, но стимулирует синтез протеогликанов в культуре мезангиальных клеток и подоцитов. Пролиферация мезангиальных клеток и матрикса наблю­ дается при IgA-нефрите (143, 165), диабетической нефропатии (102, 133, 276). Гиперпродук­ ция матрикса вследствие пролиферации клеток за вызывает гломерулосклероз.

71

F. Компоненты плазмы

59 60 61 62 63 64 65 66 67 68 69

72 73 74 75 76

Компоненты плазмы влияют на гломерулярный фильтр не новая (164), в том числе на прони­ цаемость белка (34, 98). Низкая концентрация альбумина в плазме повышает проницаемость­ гломерулярного фильтра для воды (63, 191) и макромолекул (92). Белок острой фазы воспале­ ния орсомукоид также повышает проницаемость капилляров почек для макромолекул (58, 114, 115, 117, 139). Однако механизм этого феномена неясен (168, 279, 299, 300).

81

Подоциты влияют на свойства эндотелия через секрецию сосудистого эндотелиального фак­ тора роста (VEGF) и ANG I. Гликокаликс состоит из мембранных протеогликанов синдикана и глипикана. Эндотелиальный поверхностный слой состоит из секретируемых протеоглика­ нов (версикан, перлекан), гиулорана и адсорбированных плазменных белков (орсомукоид, альбумин).

82

III. Методы изучения гломерулярного фильтра

77 78 79 80

83 84 85

Проницаемость гломерулярного фильтра изучается с помощью анализа мочи, микропункции отдельных нефронов, изолированной перфузии почек, иммуногистохимии, изоляции гломе­ рулы, гломерулярных клеток, базальной мембраны и создания искусственных мембран. Стр. 2 из 22

86 87 88

IV. Растворы, используемые для изучения гломерулярной проницаемости Проницаемость гломерулярного фильтра изучается с помощью теоретически идеальных растворов, белка, полимеров декстрана и фиколла.

90

V. Свойства растворенных веществ, влияющие на гломерулярный транспорт

91

A. Молекулярный размер

89

92 93 94 95 96 97 98 99

Эквивалентный радиус маленьких пор составляет 45—50 Å (26, 212, 213), радиус больших пор — 80—100 Å (176, 212, 213, 310). Для нейтральных растворенных веществ коэффициент проницаемости уменьшается по мере увеличения размеров молекулы, что описывается четырьмя транспортными теориями: теори­ ей двух пор, распределения нормальных пор, теорией (нейтральной) фибриллярной матрицы, теорией отрицательно заряженных волокон (136, 137). Теория отрицательно заряженных во­ локон, включает влияние заряда и размера вещества, что соотносится с биологическими дан­ ными.

100

B. Заряд молекулы

101

Независимо от теоретических моделей, заряд молекулы влияет на ее транспорт.

102

C. Конфигурация

106

Конфигурация существенно влияет на транспорт растворенных веществ через гломеруляр­ ный фильтр, если растворенные вещества имеют удлиненную или случайную форму (71, 73, 214, 247). Однако, небольшие отклонения от идеальной сферической формы существенно не влияют на транспорт (28).

107

D. Относительное значение селективности по отношению к заряду и размеру

103 104 105

111

Чтобы предсказать трансгломерулярный транспорт растворенных веществ, необходимо знать его молекулярный размер (константу диффузии, например, радиус Стокса-Эйнштейна), его заряд (плотность заряда) и конфигурацию (214). Для растворенных веществ, не имеющих случайной или удлиненной формы, она существенно не влияет на транспорт (28).

112

VI. Физические принципы и теоретические модели

108 109 110

113 114

A. Ограничение транспорта и селективность мембраны относительно размера

120

Определяющей чертой ультрафильтрационной мембраны является то, что она может выпол­ нять роль сита для макромолекул на основе их молекулярных размеров. Растворенные веще­ ства непрофильтровавшиеся имеют молекулярную массу ~1—1000 кДа. Однако, линейные размеры молекул имеют прямое отношение к их способности проходить через небольшие поры. Наиболее распространенной из таких мер молекулярного размера является радиус Стокса-Эйнштейна.

121

B. Проницаемость однородных мембран

115 116 117 118 119

122 123

Экспериментальной мерой мембранной селективности при ультрафильтрации является коэф­ фициент проницаемости. Стр. 3 из 22

124

C. Влияние молекулярной формы, размера и заряда на селективность мембран

125

1. Влияние заряда

129

Эффекты размера и заряда не совсем отделимы друг от друга. Ограниченность попыток пред­ сказать селективность относительно размера состоит в том, что фактическая структура мем­ браны может быть гораздо более сложная, чем имеющиеся модели (например, однородной цилиндрической поры или случайно ориентированные волокна) (52, 122, 134).

130

2. Влияние формы

126 127 128

135

В гелеподобной структуре эндотелиального гликокаликса и ГБМ, а также физиологического значения глобулярных белков, разделение жестких частиц в фибриллярной матрице является особенно актуальным. В таких системах, влияние молекулярной формы, похоже, имеет не­ большое значение, если растворенные вещества заметно удлиненные или они случайной формы.

136

D. Проницаемость в многослойных мембранах

131 132 133 134

137 138 139 140

Поскольку клубочковый фильтрационный барьер состоит из трех слоев (фенестрированный эндотелий с гликокаликсом, ГБМ и эпителиальные фильтрационные щели с щелевыми диафрагмами), то проницаемость в многослойной мембране отличается от таковой, в одно­ слойной.

144

Взаимозависимость коэффициентов проницаемости слоев проистекает из двух физических ограничений. Один заключается в том, что поток воды и растворенных веществ через каж­ дый слой должен быть одинаковым. Другой сдерживающий механизм заключается в том, что концентрация на нижнем слое связана с таковой на верхней стороне слоя.

145

VII. Механизмы развития протеинурии и нефротических синдромов

146

A. Протеинурия

141 142 143

149

Протеинурия — состояние, когда в моче содержится более 300 мг белка в сутки (или 200 мг/л). При нефротическом синдроме протеинурия составляет 3,5 г/сут. Микроальбумину­ рия — 30—300 мг/сут (20—200 мг/л) (31, 111, 119, 128, 183, 336).

150

B. Источник белка в моче

147 148

152

1) Фильтрация белка; 2) канальцевая секреция белков из крови; 3) синтез белка клетками по­ чек; секреция белка мочевыделительной системой и простатой (84).

153

C. Гипотеза восстановления альбумина

151

156

Протеинурия при диабетической нефропатии (226, 229) или других почечных заболеваниях (265), отражает дефекты канальцевой системы при ненарушенной клубочковой проницаемо­ сти (265, 266).

157

D. Заболевания человека

154 155

158 159 160 161 162 163

Протеинурия является отличительной чертой гломерулонефрита, диабетической нефропатии и многих других заболеваний. Среди условий, которые вызывают нефротический синдром (протеинурия, отеки, низкой концентрацией сывороточного альбумина и гиперлипидемия), минимальным изменением является нефроз. Пациенты с нефротическим синдромом финского типа имеют дефекты в гене нефрина, NPHS1 (153). Стероидоустойчивый нефротический синдром у детей обусловлен дефектом Стр. 4 из 22

164 165

гена NPHS2 (29). Ген LMX1B дефектен у больных с пателлярным синдромом, обусловлен­ ным нарушением дифференцировки подоцитов (192).

170

Протеинурия может также развиваться при ишемии-реперфузии (9, 294). Длительный период ишемии (60 мин) вызывает неселективное повреждение гломерулярного фильтра (252). При ишемия в течение 15—20 мин, последующая реперфузия главным образом влияет на гломе­ рулярную селективность относительно заряда (6, 252). Эти изменения происходили без по­ вреждения подоцитов, ГБМ и эндотелиальных клеток (6).

171

Литература

166 167 168 169

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