A long-tailed pair as an input stage... with a single-ended output and, regretfully, without a dynamic load in the cathodes... a resisitve "shifting down" coupling to the second stage (the left section of the second tube)...
The two neon bulbs in series "shift down" the anode voltage variations... then a cathode follower boosts the output... These 680 k and 4.7 M resistors in series to the neon bulbs are a bit confusing... but obviously, the grid current is negligible... like today FETs... The 7.5 pF bypass capacitor shunts (for AC) the high ohmic "moving" network.
But what is the role of the 221 kom resistor connecting the cathodes of the second tube? It "smacks" of a positive feedback increasing the cathode resistor of 9.1 k - another famous idea (cathode degeneration). The 500 pF bypass capacitor shunts (for AC) the relatively high ohmic cathode resistor. Or maybe, the two sections of the second tube form another long-tailed pair? But it seems unlikely...
Another great idea is the ±300 V split supply...
I think there is no more ideas for unraveling... Of course, I could browse through Google to find some explanations... but it is so exciting to find yourself the new idea... especially when you do not know well tubes... and you have to convey basic ideas from transistors and semiconductor op-amps...
In this connection, maybe it would be interesting to compare it with the 741 op-amp internal structure? Oh my God ! Only 20 transistors versus 2 tubes! What a perfect simplicity!
By my opinion this resistor defines the positive feedback (DC) - 500 pF interrupts the feedback for higher frequencies. This solution gives us a high value of DC gain.
It seems it "interrupts" both the negative feedback introduced by the 9.1 k resistor (the cathode degeneration) and the positive feedback introduced by the 221 k resistor?