Where and why were capital letters first used in English headlines?

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In the medieval period there were two basic styles of writing: Roman and Irish. The Roman writing style was those big capitals you see on monuments in museums like this:

M·AGRIPPA·L·F·COS·TERTIVM·FECIT

The Irish style was developed in Irish monasteries and spread throughout Europe as those monks moved around (see the book, "How the Irish Saved Civilization" for more info). This style is characterized by what we call a "lower case" letter, with very large ornamental letters at the beginning of chapters.

Roman writing was used for civil purposes: book keeping, laws, property records. Irish writing was used for literature, religious writing and scholarship.

When printing developed at the end of 15th century what happened was that there was a merger of these two styles. Roman letters became "capitals" and Irish letters became "lower case". Case refers to the boxes with printing type as it is kept at a desk. As litlnemo says, there was no set method for combining the two styles. Different printers did whatever seemed most readable. Over time our current practice eventually evolved.

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This discussion on the Linguist List discusses the frequent use of capitalized words in early printed materials, which was used to mark salience -- importance. The comment says:

"You'll find it to varying degrees in most of the major languages of the region during this time period... In the English-speaking world, the practice was maintained, at least to a certain degree, well into the 18th century. Since then, it has been more and more restricted to newspaper headines (hence the common label `headline capitalization'), and even there it's been disappearing in recent decades."

A question on english.stackexchange.com discusses the capitalization of nouns in English in the 17th-18th centuries, but not specifically about titles.

The thing is, looking at early printed titles, it is difficult to tell whether they are using caps because of some attempt at a sort of "title case," or it's that they would have used the caps anyway because that was the way they tended to write. For example, what would you make of this one, from 1579?

Sermons of John Calvin, 1579

It isn't really what we would call title case, but it does resemble it.

This one from 1613 only seems to capitalize a few emphasized words -- Wonders, Sea, Land, World.

enter image description here

Going through a pile of images of early books and some Medieval manuscripts, I'm finding that there really isn't much consistency. Words started to be capitalized for emphasis, but you see this even as early as the late 14th-early 15th centuries (in at least one manuscript of the cookbook Forme of Cury, for example -- which uses "title case" in a few headlines but not in most others.) And it seems to vary like crazy over the next couple of centuries. My opinion is that the use of caps to mark salience gradually developed into a slightly more standardized use of them to mark titles. It doesn't seem to have specifically come from German.

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