In the Data Encryption Standard (DES), S-boxes (Substitution boxes) are the core elements that provide non-linearity and security. During each of the 16 Feistel rounds, the 32-bit right half of the data is expanded to 48 bits, XORed with the round key, and divided into eight 6-bit blocks. Each block is then passed through one of eight fixed S-boxes. An S-box maps 6 input bits to 4 output bits using a lookup table, reducing 48 bits to 32 bits. This substitution introduces confusion, making the relationship between key and ciphertext complex, and is vital for DES security.