Frontispiece · Ordo Filtrantia

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Frontispiece · ORD. FILTRANTIA

Ordo Filtrantia

the filters · the urinary system

The plates

The eight specimens

Four families, and the split between them is the lesson. One plate for the organ, four for the nephron that actually makes the urine, one for the collecting system that is not part of the nephron at all, and two for the road out, where nothing is changed any more.

Pl. I

Ren zonatus var. actinidius

the sorting house · kidney

A kiwifruit cut across, drawn in coloured pencil on a transparent ground: a brown fuzzy skin, a pale core sending rays outward, a ring of small black seeds, and green flesh between.

Open Pl. I →

Pl. II

Corpusculum fenestratum var. cyathi

the sieve in a cup · renal corpuscle

A bird's nest fungus drawn in coloured pencil on a transparent ground: a fluted brown cup holding a loose cluster of pale discs, with a clear gap all the way around them.

Open Pl. II →

Pl. III

Tubulus villosus var. callistemonis

the greedy one · proximal convoluted tubule

A bottlebrush flower spike drawn in coloured pencil on a transparent ground: hundreds of fine rust-red filaments radiating from a woody stem, with narrow leaves below.

Open Pl. III →

Pl. IV

Tubulus tenuis var. ardeae

the long game · nephron loop, thin limb

A great blue heron drawn in coloured pencil on a transparent ground: slate blue-grey, neck folded back, long dagger bill, one leg standing and one lifted and folded.

Open Pl. IV →

Pl. V

Tubulus densus var. mimosae

the sensor · distal tubule and macula densa

A sensitive plant drawn in coloured pencil on a transparent ground: a stem carrying one compound leaf fully open, one folded shut, and one halfway closed between them.

Open Pl. V →

Pl. VI

Ductus discretus var. sphagni

the gatekeeper · collecting duct

A sphagnum moss shoot drawn in coloured pencil on a transparent ground: a dense head above whorls of spreading and hanging branches.

Open Pl. VI →

Pl. VII

Via stellata var. carambolae

the star · ureter

A starfruit drawn in coloured pencil on a transparent ground, with one end cut off so the five-pointed star of the cut face shows and the ridges run back along the body.

Open Pl. VII →

Pl. VIII

Via trilaminata var. octopodis

the squeeze · bladder

A common octopus drawn in coloured pencil on a transparent ground: a rounded rust-coloured mantle above the head, and eight arms curling outward with suckers along them.

Open Pl. VIII →

The ground plan

One tube, twice as long as it looks

Every nephron is a single unbranched tube with a filter bolted to the front of it, and almost all of its length is spent taking back what the filter threw away. About 180 litres of plasma are filtered a day and about 1.5 litres leave as urine, so roughly 99 % of what crosses the filter is reclaimed further down. Filtering happens only in the cortex. Concentrating happens only in the medulla. Nearly everything else on this page follows from those two facts.

Fig. 1 · one nephron, end to end

Hand-drawn figure in coloured pencil: one nephron drawn end to end across a cortex above and a medulla below, the boundary marked by a horizontal rule. In the cortex, a renal corpuscle with its afferent and efferent arterioles, a convoluted proximal tubule drawn with a fuzzy brush border, and a distal tubule returning to touch its own corpuscle at a patch of crowded nuclei. The nephron loop descends into the medulla and turns back. The collecting duct is drawn in a different colour, slate blue, running straight down to the papilla, with two other nephrons joining it — because the collecting duct is not part of the nephron.

Filter first, then three tubules that each take back something different — and the collecting duct in slate blue, because it belongs to a different family.

The five words

  • Cortex — the grainy outer zone. Every renal corpuscle in the kidney sits here, which is why filtration happens only here.
  • Medulla — the striped inner zone, arranged in pyramids. Everything in it is cut lengthwise — loops, collecting ducts and straight vessels, all running the same way.
  • Nephron — one corpuscle and its tubules, from the filter to the point where it joins a collecting duct. A million per kidney, and the whole of urine production happens inside one.
  • Collecting system — the ducts the nephrons drain into. Not part of the nephron: different embryonic origin, different job, and the only place a hormone finishes the work.
  • Filtrate — what crosses the filter. It is not urine yet, and it will not be until the last duct has had its say.

Decide whether you are in cortex or medulla before anything else. It settles half the question on its own.

First question

Where are you?

Before you look at a single cell, work out which zone of the kidney you are in — or whether you have left the kidney altogether.

Fig. 2 · where are you

Hand-drawn figure in coloured pencil: three framed microscope fields side by side. On the left, cortex — four renal corpuscles among round tubule profiles. In the middle, medulla — eleven tubules all cut lengthwise and all running the same way, one of them drawn in slate blue as a collecting duct. On the right, outside the kidney — a ureter in cross-section with a star-shaped lumen inside a thick muscular wall.

Three fields. Corpuscles, then everything cut lengthwise, then a wall thick enough to be a road.

  • Cortex — round corpuscles scattered among tubules cut every which way. One corpuscle anywhere in the field is enough to settle it.
  • Medulla — no corpuscles at all, and everything running parallel because it is all cut lengthwise.
  • Outside the kidney — a lumen with a thick muscular wall around it and domed cells on the surface. That is the road out, not the kidney.

Corpuscles are the test. Their absence is not proof on its own, because a badly cut cortical field can miss them — so ask whether the tubules are running parallel as well.

Second question

What is the lining doing?

Filtration is indiscriminate: it throws out everything small enough to go. Every tubule after it is choosy, and the lining tells you how choosy.

Fig. 3 · what the lining is doing

Hand-drawn figure in coloured pencil: four strips of epithelium side by side, numbered 1 to 4. One, tall cells with a brush border of microvilli standing on the surface — a proximal tubule, taking back nearly everything. Two, shorter cells with a clean open lumen — a distal tubule. Three, boxy pale cells with sharply drawn borders between them — a collecting duct. Four, urothelium: domed umbrella cells on top over two further layers of rounded cells — ureter and bladder. A line beneath reads, the lining tells you how choosy the tube is.

Four linings, from taking back nearly everything to taking back nothing at all.

  • Brush-bordered — fuzz filling the lumen so you cannot find its edge. Taking back nearly everything: the proximal convoluted tubule.
  • Bare cuboidal — cube-shaped cells, a clean open lumen, nothing on the free surface. Taking back only sodium, and only on orders: the distal convoluted tubule.
  • Sharp-bordered — pale boxy cells with knife-edged joins between them. Taking back water only when a hormone says so: the collecting duct.
  • Urothelium — several rows of cells with big domed ones on top. Taking back nothing, ever — which is exactly why it is waterproof: ureter and bladder.

A fuzzy lumen beats everything else in the kidney. If you cannot find the edge of the lumen, you have a proximal tubule and nothing more needs deciding.

The naming key

How to read a name

Two words. The first says where you are in the path; the second says what the lining is doing there.

First word · where you are

  • Ren — the whole organ
  • Corpusculum — the filter itself
  • Tubulus — a tubule of the nephron
  • Ductus — the collecting system, which is not the nephron
  • Via — the road out

Second word · what the lining is doing

  • zonatus — zoned — cortex, medulla, pelvis
  • fenestratum — windowed — holes straight through
  • villosus — shaggy — a brush border
  • tenuis — one cell thin, and nothing on it
  • densus — crowded — cells packed close and sensing
  • discretus — sharp — knife-edged borders
  • stellata — folded to a star
  • trilaminata — three coats of muscle

Worked both ways

  • Tubulus villosus → a nephron tubule with a brush border → the proximal convoluted tubule.
  • Via stellata → the road out, with a star-shaped lumen → the ureter.
  • You saw pale boxy cells with knife-edged borders running through the medulla → Ductus discretus → a collecting duct.

Three that trip people up

The look-alikes

Three pairs account for most of the mistakes in this volume. Each one has a single reliable test, and not one of those tests is size.

Fig. 4 · three that trip people up

Hand-drawn figure in coloured pencil: three look-alike pairs stacked. Top, a proximal tubule whose lumen is filled with brush border beside a distal tubule with a clean open lumen and more nuclei around it. Middle, a thin limb of the nephron loop, empty, beside a vas rectum holding red blood cells. Bottom, a ureter with a star-shaped lumen beside a length of bladder wall showing folded urothelium over three muscle coats.

Three pairs, stacked. Lumen, then contents, then wall.

A proximal tubule, or a distal one?

The trap: both are cuboidal, and both sit in the cortex.

They are neighbours in the same field and much the same shape, so position will not separate them. The proximal tubule carries a brush border that fills its lumen with fuzz and makes the edge impossible to find. The distal tubule has a clean open lumen you could draw a line around. The proximal one is also the wider of the two, with fewer nuclei in a cross-section, because its cells are bigger.

Look for the edge of the lumen. If you can find it, it is distal.

A thin limb, or a vas rectum?

The trap: both are thin-walled tubes running lengthwise through the medulla.

Both are one flat cell thick and both run with the pyramids, so neither shape nor direction is any help. The vas rectum is a blood vessel, so there are red blood cells sitting in it. The thin limb is a tubule, so it is empty, or holds nothing but pale filtrate.

Check the contents, not the wall. Red cells inside means a vessel.

A ureter, or a bladder?

The trap: both are lined with urothelium.

The lining is identical in the two, so it cannot separate them. The ureter has a lumen folded into a star, and nothing else in the body has that outline; its wall carries two coats of muscle. The bladder has a wide, shallow lumen and three coats that cross one another, which is what lets it squeeze itself empty from every direction at once.

A star means ureter. Count the muscle coats to confirm it.

Keying out a specimen

Four questions, in this order

This is the procedure. It works on a urinary slide you have never seen before and have not been told the name of, which is the only test that matters.

  1. Are you inside the kidney at all? — A thick muscular wall around the lumen, with domed cells on the surface, means you have left it — go to Pl. VII or Pl. VIII.
  2. Cortex or medulla? — One corpuscle anywhere in the field means cortex. No corpuscles, and everything cut lengthwise, means medulla.
  3. Can you see the lumen clearly? — Full of fuzz, with no findable edge, means a proximal convoluted tubule. Clean and open means a distal tubule or a collecting duct.
  4. Are the cell borders sharp? — Knife-edged, boxy and pale means a collecting duct, wherever in the kidney you happen to be.

Three overrides. A fuzzy lumen beats everything — a brush border is the single most reliable mark in the kidney. Sharp borders beat location, because collecting ducts run through cortex and medulla alike and look the same in both. Domed surface cells outside the kidney mean urothelium — and then the lumen decides which, because a star is only ever a ureter.

YOur missions

Field Notes for Future Missions

Before you go to the plates, try the key on the whole path at once.

Mission 1 · follow the filtrate down

Mission 2 · key it out

Field notes

Why eight organisms

A kiwifruit, a bird’s nest fungus, a bottlebrush flower, a great blue heron, a sensitive plant, a cushion of sphagnum moss, a starfruit and an octopus have nothing to do with one another and nothing to do with kidneys. That is rather the point of them.

A list of eight structures is something you memorise and then lose. Eight organisms that each carry one structural idea — drainage running outward through productive tissue, a sieve sitting in a cup, a surface multiplied into fuzz, a long thin counterflow, a patch that senses and reports, a gate that opens only when told, a star-shaped lumen, three crossing coats of muscle — stay put, because you can picture them. The name is a handle for the picture, not the other way round.

Begin · Pl. I

Ren zonatus var. actinidius

the sorting house · kidney.

List of terms