Makers of British botany/Stephen Hales 1677—1761

by Francis Darwin

Plate VIII

STEPHEN HALES (1759)

STEPHEN HALES
1677—1761

By FRANCIS DARWIN

An error corrected—Hales' scientific contemporaries—Physiology or Physics—Hales the Founder of the experimental method in Physiology—His style—Cambridge days—Teddington—Vegetable Staticks—Experiments described—Transpiration—Root Pressure—Assimilation—Practical application to greenhouses—Distribution of growth first measured—Hales' other activities—Sachs' tribute.

In attempting to give a picture of any man's life and work it is well to follow the rule of the Dictionary of National Biography, and begin with the dates of his birth and death. Stephen Hales was born in 1677 and died in 1761, having had experiences of the reigns of seven sovereigns.

The authorities for the life of Hales are given in my article on him in the Dictionary of National Biography. Botanists in general probably take their knowledge of the main facts of his life from Sachs' History of Botany. It is therefore worth while to point out that both the original and the English translation (1890) contain the incorrect statement that Hales was educated at Christ's College, Cambridge, and that he held the living of Riddington, whereas he is one of the glories of Corpus, and was perpetual curate of Teddington. These inaccuracies however are trifles in relation to the great and striking merits of Sachs' History, a work which to my thinking exhibits the strength and brilliance of the author's mind as clearly as any of his more technical writings. Sachs was no niggling biographer, and his broad vigorous outlines must form the basis of what anyone, who follows him, has to say about the Botanists of a past day.

To return to Hales' birth: it is of interest to note how he fits into the changing procession of lives, to see what great men overlap his youth, who were his contemporaries in his maturity, and who were appearing on the scientific stage as he was leaving it.

Sir Isaac Newton was the dominant figure in English science while Hales was developing. He died in 1727, the year in which Hales published his Vegetable Staticks, a book, which like the Origin of Species, appeared when its author was 50 years of age; Newton was at the zenith of his fame when Hales was a little boy of 10—his Principia having been published in 1687. And when Hales went up to Cambridge in 1696 he must have seen the great man coming from his rooms^[1] in the N.E. corner of the Great Court of Trinity—that corner where Newton's and other more modern ghosts surely walk—Macaulay who used to read, pacing to and fro by the chapel^[2], and Thackeray who, like his own Esmond, lived "near to the famous Mr Newton's lodgings." In any case there can be no doubt that the genius of Newton cast its light on Hales, as Sachs has clearly pointed out (Hist. Bot., Eng. Tr., p. 477). Another great man who influenced Hales was Robert Boyle, who was born 1627 and died 1691. John Mayow again, that brilliant son of Oxford, whose premature death at 39 in 1679 was so heavy a blow to science, belongs to the same school as Hales—the school which was within an ace of founding a rational chemistry, but which was separated from the more obvious founders of that science by the phlogiston-theory of Becchers and Stahl. I do not find any evidence that Hales was influenced by the phlogistic writers and this is comprehensible enough, if, as I think, he belongs to the school of Mayow and Boyle.

The later discoverers in chemistry are of the following dates, Black 1728—1799, Cavendish 1731—1810, Priestley 1733—1804, Scheele 1742—1786, Lavoisier 1743, guillotined 1794. These were all born about the time of Hales' zenith, nor did he live^[3] to see the great results they accomplished. But it should not be forgotten that Hales' chemical work made more easy the triumphant road they trod.

I have spoken of Hales in relation to chemists and physicists because, though essentially a physiologist, he seems to me to have been a chemist and physicist who turned his knowledge to the study of life, rather than a physiologist who had some chemical knowledge.

Whewell points out in his History of the Inductive Sciences^[4] that the Physiologist asks questions of Nature in a sense differing from that of the Physicist. The Why? of the Physicist meant Through what causes? that of the Physiologist—To what end? This distinction no longer holds good, and if it is to be applied to Hales it is a test which shows him to be a physicist. For, as Sachs shows, though Hales was necessarily a teleologist in the theological sense, he always asked for purely mechanical explanations. He was the most unvitalistic of physiologists, and I think his explanations suffered from this cause. For instance, he seems to have held that to compare the effect of heat on a growing root to the action of the same cause on a thermometer^[5] was a quite satisfactory proceeding. And there are many other passages in Vegetable Staticks where one feels that his speculations are too heavy for his knowledge.

Something must be said of Hales' relation to his predecessors and successors in Botanical work. The most striking of his immediate predecessors were Malpighi 1628—1694, Grew 1628—1711, Ray 1627—1705, and Mariotte (birth unknown, died 1684); and of these the three first were born one hundred years before the publication of Vegetable Staticks. Malpighi and Grew were essentially plant-anatomists, though both dealt in physiological speculations. Their works were known to Hales, but they do not seem to have influenced him.

We have seen that as a chemist Hales is somewhat of a solitary figure, standing between what may be called the periods of Boyle and of Cavendish. This is even more striking in his Botanical position, for here he stands in the solitude of all great original inquirers. We must go back to Van Helmont, 1577—1644, to find anyone comparable to him as an experimentalist. His successors have discovered much that was hidden from him, but consciously or unconsciously they have all learned from him the true method and spirit of physiological work.

It may be urged that in exalting Hales I am unfair to Malpighi. It may be fairer to follow Sachs in linking these great men together and to insist on the wonderful fact that before Malpighi's book in 1671, vegetable physiology was still where Aristotle left it, whereas 56 years later in 1727 we find in Hales' book an experimental science in the modern sense.

It should not be forgotten that students of animal physiology agree with botanists as to Hales' greatness. A writer in the Encyclopædia Britannica speaks of him as "the true founder of the modern experimental method in physiology."

According to Sachs, Ray made some interesting observations on the transmission of water, but on the whole what he says on this subject is not important. There is no evidence that he influenced Hales.

Mariotte the physicist came to one physiological conclusion of great weight^[6]; namely, that the different qualities of plants, e.g. taste, odour, etc., do not depend on the absorption from the soil of differently scented or flavoured principles, as the Aristotelians imagined, but on specific differences in the way in which different plants deal with identical food material—an idea which is at the root of a sane physiological outlook. These views were published in 1679^[7], and may have been known to Hales. He certainly was interested in such ideas, as is indicated by his attempts to give flavour to fruit by supplying them with medicated fluids. He probably did not expect success for he remarks, p. 360: "The specifick differences of vegetables, which are all sustained and grow from the same nourishment, is [sic] doubtless owing to the very different formation of their minute vessels, whereby an almost infinite variety of combinations of the common principles of vegetables is made." He continues in the following delightful passage: "And could our eyes attain to a sight of the admirable texture of the parts on which the specific differences in plants depends [sic] what an amazing and beautiful scene of inimitable embroidery should we behold? what a variety of masterly strokes of machinery? what evident marks of consummate wisdom should we be entertained with?" To conclude what has been said on Hales' chronological position—Ingenhousz, the chief founder of the modern point of view on plant nutrition, was born 1730 and published his book On Vegetables, etc. in 1779. So that what was said of Hales' chemical position is again true of him considered in relation to nutrition; he did not live to see the great discoveries made at the close of the 18th century.

There is in his writing a limped truthfulness and simplicity, unconsciously decorated with pretty 18th century words and half-rusticities which give it a perennial charm. And inasmuch as I desire to represent Hales not merely as a man to be respected but also to be loved, it will be as well to give what is known of the personal side of his character before going on to a detailed account of his work.

He was, as we have seen, entered at Corpus Christi College, Cambridge, in June, 1696. In February, 1702–3, he was admitted a fellow of the College. It was during his life as a fellow that he began to work at chemistry in what he calls "the elaboratory in Trinity College." The room is now occupied by the Senior Bursar and forms part of the beautiful range of buildings in the bowling green, which, freed from stucco and other desecration, are made visible in their ancient guise by the piety of a son of Trinity and the wisdom of the College authorities. It was here, according to Dr Bentley, that "the thieving Bursars of the old set embezzled the College timber^[8]," and it was this room that was fitted up as "an elegant laboratory" in 1706 for John Francis Vigani, an Italian chemist, who had taught unofficially in the University for some years and became the first Professor of Chemistry at Cambridge in 1703.

Judging from his book, Medulla Chymiae, 1682, Vigani was an eminently practical person who cared greatly about the proper make of a furnace and the form of a retort, but was not cumbered with theories.

Hales vacated his fellowship and became minister or perpetual curate of Teddington^[9] in 1708—9 and there he lived until his death, fifty-two years afterwards. He was married (?1719) and his wife died without issue in 1721.

He attracted the attention of Royalty, and received plants from the King's garden at Hampton Court. Frederick Prince of Wales, the father of George III, is said to have been fond of surprising him in his laboratory at Teddington. This must surely be a unique habit in a prince, but we may remember that, in the words of the Prince's mock epitaph, "since it is only Fred there's no more to be said." He became Clerk of the Closet to the Dowager Princess and this "mother of the best of Kings" as she calls herself put up his monument in Westminster Abbey. Hales had the honour of receiving the Copley Medal from the Royal Society in 1739, and Oxford made him a D.D. in 1733.

Some years ago I made a pilgrimage to Teddington and found, in the parish registers, many interesting entries by his hand; the last in a tremulous writing is on November 4th, 1760, two months before he died. He was clearly an active parish priest. He made his female parishioners do public penance when he thought they deserved it: he did much for the fabric of the church. "In 1754^[10] he helped the parish to a decent water supply and characteristically records, in the parish register, that the outflow was such as to fill a two-quart vessel in 'three swings of a pendulum beating seconds, which pendulum was 39 + ²⁄₁₀ inches long from the suspending nail to the middle of the plumbet or bob'." Under the tower he helped to build (which now serves as a porch) Stephen Hales is buried, and the stone which covers his body is being worn away by the feet of the faithful. By the piety of a few botanists a mural tablet, on which the epitaph is restored, has been placed near the grave.

Horace Walpole called Hales "a poor, good, primitive creature" and Pope^[11] (who was his neighbour) said "I shall be very glad to see Dr Hales, and always love to see him, he is so worthy and good a man." Peter Collinson writes of "his constant serenity and cheerfulness of mind"; it is also recorded that "he could look even upon wicked men, and those who did him unkind offices, without any emotion of particular indignation; not from want of discernment or sensibility; but he used to consider them only like those experiments which, upon trial, he found could never be applied to any useful purpose, and which he therefore calmly and dispassionately laid aside."

Hales' work may be divided into three heads:

I.	Physiological, animal and vegetable;
II.	Chemical;
III.	Inventions and miscellaneous essays.

Under No. I. I shall deal only with his work on plants. The last heading (No. III.) I shall only refer to slightly, but the variety and ingenuity of his miscellaneous publications is perhaps worth mention here as an indication of the quality of his mind. It seems to me to have had something in common with the versatile ingenuity of Erasmus Darwin and of his grandson Francis Galton. The miscellaneous work also exhibits Hales as a philanthropist, who cared passionately for bettering the health and comfort of his fellow creatures by improving their conditions of life.

His chief book from the physiological and chemical point of view is his Vegetable Staticks. It will be convenient to begin with the physiological part of this book, and refer to the chemistry later. Vegetable Staticks is a small 8vo of 376 pages, dated on the title-page 1727. The "Imprimatur Isaac Newton Pr. Reg. Soc." is dated February 16, 172⁶⁄₇, and this date is of some slight interest, for Newton died on March 20, and Vegetable Staticks must have been one of the last books he signed.

The dedication is to George Prince of Wales, afterwards George III. The author cannot quite avoid the style of his day, for instance: "And as Solomon the greatest and wisest of men, deigned^[12] to inquire into the nature of Plants, from the Cedar of Lebanon, to the Hyssop that springeth out of the wall. So it will not, I presume, be an unacceptable entertainment to your Royal Highness," etc.

But the real interest of the dedication is its clear statement of his views on the nutrition of plants. He asserts that plants obtain nourishment, not only from the earth, "but also more sublimed and exalted food from the air, that wonderful fluid, which is of such importance to the life of Vegetables and Animals," etc. We shall see that his later statement is not so definite, and it is well to rescue this downright assertion from oblivion.

His book begins with the research for which he is best known, namely that on transpiration. He took a sunflower growing in a flower-pot, covering the surface of the earth with a plate of thin milled lead, and cemented it so that no vapour could pass, leaving a corked hole to allow of the plant being watered. He did not take steps to prevent loss through the pot, but at the end of the experiment cut off the plant, cemented the stump and found that the "unglazed porous pot" perspired 2 ozs. in 12 hours, and for this he made due allowance.

The plant so prepared he proceeded to weigh at stated intervals. He obtained the area of the leaves by dividing them into parcels according to their several sizes and measuring one leaf^[13] of each parcel. The loss of water in 12 hours converted to the metric system is 1.3 c.c. per 100 sq. cm. of leaf-surface; and this is of the same order of magnitude as Sachs' result^[14], namely 2.2 c.c. per 100 sq. cm.

He goes on to measure the surface of the roots^[15] and to estimate the rate of absorption per area. The calculation is of no value, since he did not know how small a part of the roots is absorbent, nor how enormously the surface of that part is increased by the presence of root-hairs. He goes on to estimate the rate of the flow of water up the stem; this would be 34 cubic inches in 12 hours if the stem (which was one square inch in section) were a hollow tube. He then allowed a sunflower stem to wither and to become completely dry, and found that it had lost ³⁄₄ of its weight, and assuming that the ¹⁄₄ of the "solid parts" left was useless for the transmission of water he increases his 34 by ¹⁄₃ and gives 45+¹⁄₃ cubic inches in 12 hours as the rate. But the solid matter which he neglected contained the vessels and he would have been nearer to the truth had he corrected his figures on this basis. The simplest plan is to compare his results with those obtained by Sachs^[16] in allowing plants to absorb solutions of lithium-salts. If the flow takes place through conduits equivalent to a quarter of a square inch in area, the fluid will rise in 12 hours to a height of 4 x 34 or 136 inches or in one hour to 28.3 cm.^[17] This is a result comparable to, though very much smaller than, Sachs' result with the sunflower, viz. 63 cm. per hour.

The data are however hardly worth treating in this manner. But it is of historic interest to note that when Sachs was at work on his Pflanzenphysiologie, published in 1865, he was compelled to go back nearly 140 years to find any results with which he could compare his own.

We need not follow Hales into his comparison between the "perspiration" of the sunflower and that of a man, nor into his other transpiration experiments on the cabbage, vine, apple, etc. But one or two points must be noted. He found^[18] the "middle rate of perspiration" of a sunflower in 12 hours of daylight to be 20 ounces, and that of a "dry warm night" about 3 ounces; thus the day transpiration was roughly seven times the nocturnal rate. This difference may be accounted for by the closure of the stomata at night.

Hales of course knew nothing of stomata, but it is surprising to find Sachs in 1865 discussing the problem of transpiration with hardly a reference to the effect of stomatal closure.

Hales^[19] notes another point which a knowledge of stomatal behaviour might have explained, viz. that with "scanty watering the perspiration much abated," he does not attempt an explanation but merely refers to it as a "healthy latitude of perspiration in this Sunflower."

In the course of his work on sunflowers he notices that the flower follows the sun, he says however that it is "not by turning round with the sun," i.e. that it is not a twisting of the stalk, and goes on to call it nutation which must be the locus classicus for the term used in this sense.

An experiment^[20] that I do not remember to have seen quoted elsewhere is worth describing. It is one of the many experiments that show the generous scale on which his work was planned. An apple bough five feet long was fixed to a vertical glass tube nine feet long. The tube being above and the branch hanging below the pressure of the column of water would act in concert with the suck of the transpiring leaves instead of in opposition to this force. He then cut the bare stem of his branch in two, placing the apical half of the specimen (bearing side branches and leaves) with its cut end in a glass vessel of water, the basal and leafless half of the branch remained attached to the vertical tube of water. In the next 30 hours only 6 ounces dripped through the leafless branch, whereas the leafy branch absorbed 18 ounces. This, as he says, shows the great power of perspiration. And though he does not pursue the experiment, it is worthy of note as an attempt like those of Janse^[21] and others to correlate the flow of water under pressure with the flow due to transpiration.

It is interesting to find that Hales used the three methods of estimating transpiration which have been employed in modern times, namely, (i) weighing, (ii) a rough sort of potometer, (iii) enclosing a branch in a glass balloon and collecting the precipitated moisture, the well-known plan followed by various French observers.

He (Vegetable Staticks, p. 51) concluded his balance of loss and gain in transpiring plants by estimating the amount of available water in the soil to a depth of three feet, and calculating how long his sunflower would exist without watering. He further concludes (p. 57) that an annual rainfall (of 22 inches) is "sufficient for all the purposes of nature, in such flat countries as this about Teddington."

He constantly notes small points of interest, e.g. (p. 82) that with cut branches the water absorbed diminishes each day and that the former vigour of absorption may be partly renewed by cutting a fresh surface^[22].

He also showed (p. 89) that the transpiration current can flow perfectly well from apex to base when the apical end is immersed in water.

These are familiar facts to us, but we should realise that it is to the industry and ingenuity of Hales that we owe them. In a repetition (p. 90) of the last experiment, we have the first mention of a fact fundamentally important. He took two branches (which with a clerical touch he calls M and N) and having removed the bark from a part of the branch dipped the ends in water, N with the great end downwards, but M upside down. In this way he showed that the bark was not necessary for the absorption or transmission of water^[23]. I suspect that one branch was inverted out of respect for the hypothesis of sap-circulation. He perhaps thought that water could travel apically by the wood, but only by the bark in the opposite direction.

Later in his book (pp. 128 and 131) he gives definite arguments against the hypothesis in question.

Next in order (p. 95) comes his well-known experiment on the pressure exerted by peas increasing in size as they imbibe water. There are, however, pitfalls in this result of which Hales was unaware, and perhaps the chief interest to us now is that he considered the imbibition of the peas^[24] to be the same order of phenomenon as the absorption of water by a cut branch—notwithstanding the fact that he knew^[25] the absorption to depend largely on the leaves. It may be noticed that Sachs with his imbibitional view of water-transport may be counted a follower of Hales.

In order to ascertain "whether there was any lateral communication of the sap and sap vessels, as there is of blood in animals," Hales (p. 121) made the experiment which has been repeated in modern laboratories^[26], i.e. cutting a "gap to the pith" and another opposite to it and a few inches above. This he did on an oak branch six feet long whose basal end was placed in water. The branch continued to "perspire" for two days, but gave off only about half the amount of water transpired by a normal branch^[27]. He does not trouble himself about this difference, being satisfied of "great quantities of liquor having passed laterally by the gap."

He is interested in the fact of lateral transmission in connexion with the experiment of the suspended tree (Fig. 24, p. 126), which is dependent on the neighbours to which it is grafted for its water supply. This seems to be one of the results that convinced him that there is a distribution of food material which cannot be described as circulation of sap in the sense that was then in vogue.

Hales (p. 143) was one of the first^[28] to make the well-known experiment—the removal of a ring of bark, with the result that the edge of bark nearest the base of the branch swells and thickens in a characteristic manner. He points out that if a number of rings are made one above the other, the swelling is seen at the lower edge of each isolated piece of bark, and therefore (p. 143) the swelling must be attributed "to some other cause than the stoppage of the sap in its return downwards," because the first gap in the bark should be sufficient to check the whole of the flowing sap^[29]. He must in fact have seen that there is a redistribution of plastic material in each section of bark.

We now for the moment leave the subject of transpiration and pass on to that of root-pressure on which Hales is equally illuminating.

Figure from Vegetable Staticks showing a vine with mercury gauges in place to
demonstrate root-pressure.

His first experiment, Vegetable Staticks, p. 100, was with a vine to which he attached a vertical pipe made of three lengths of glass-tubing jointed together. His method is worth notice. He attached the stump to the manometer with a "stiff cement made of melted Beeswax and Turpentine, and bound it over with several folds of wet bladder and pack-thread." We cannot wonder that the making of water-tight connexions was a great difficulty, and we can sympathise with his belief that he could have got a column more than 21 feet high but for the leaking of the joints on several occasions. He notes the familiar fact that the vine-stump absorbed water before it began to extrude it.

He afterwards (pp. 106–7) used a mercury gauge and registered a root-pressure of 32+¹⁄₂ inches or 36 feet 5+¹⁄₃ inches of water which he proceeds to compare with his own determination of the blood-pressure of the horse (8 feet) and of other animals. Perhaps the most interesting of his root-pressure experiments was that (p. 110) in which several manometers were attached to the branches of a bleeding vine and showed a result which convinced him that "the force is not from the root only, but must proceed from some power in the stem and branches," a conclusion which some modern workers have also arrived at. The figure on page 77 is a simplified reproduction of the plate (Fig. 19) in Vegetable Staticks.

Assimilation.

Hales' belief that plants draw part of their food from the air, and again that air is the breath of life, of vegetables as well as of animals (p. 148), are based upon a series of chemical experiments performed by himself. Not being satisfied with what he knew of the relation between "air" (by which he meant gas) and the solid bodies in which he supposed gases to be fixed, he delayed the publication of Vegetable Staticks for some two years, and carried out the series of observations which are mentioned in his title-page as "An attempt to analyse the air, by a great variety of chymio-statical experiments" occupying 162 pages of his book^[30].

The theme of his inquiry he takes (Vegetable Staticks, p. 165) from "the illustrious Sir Isaac Newton," who believed that "Dense bodies by fermentation rarify into several sorts of Air; and this Air by fermentation, and sometimes without it, returns into dense bodies."

Hales' method consisted in heating a variety of substances, e.g. wheat-grains, pease, wood, hog's blood, fallow-deer's horn, oyster-shells, red-lead, gold, etc., and measuring the "air" given off from them. He also tried the effect of acid on iron filings, oyster-shells, etc. In the true spirit of experiment he began by strongly heating his retorts (one of which was a musket barrel) to make sure that no air arose from them. It is not evident to me why he continued at this subject so long. He had no means of distinguishing one gas from another, and almost the only quality noted is a want of permanence, e.g. when the CO₂ produced was dissolved by the water over which he collected it. Sir E. Thorpe^[31] points out that Hales must have prepared hydrogen, carbonic acid, carbonic oxide, sulphur dioxide, marsh gas, etc. It may, I think, be said that Hales deserved the title usually given to Priestley, viz. "the father of pneumatic^[32] chemistry."

Perhaps the most interesting experiment made by Hales is the heating of minium (red-lead) with the production of oxygen. It proves that he knew, as Boyle, Hooke and Mayow did before him, that a body gains weight in oxidation. Thus Hales remarks: "That the sulphurous and aereal particles of the fire are lodged in many of those bodies which it acts upon, and thereby considerably augments their weight, is very evident in Minium or Red Lead which is observed to increase in weight in undergoing the action of the fire. The acquired redness of the Minium indicating the addition of plenty of sulphur in the operation." He also speaks of the gas distilled from minium, and remarks "It was doubtless this quantity of air in the minium which burst the hermetically sealed glasses of the excellent Mr Boyle, when he heated the Minium contained in them by a burning glass" (p. 287).

This was the method also used by Priestley in his celebrated experiment of heating red-lead in hydrogen; whereby the metallic lead reappears and the hydrogen disappears by combining with the oxygen set free. This was expressed in the language of the day as the reconstruction of metallic lead by the addition of phlogiston (the hydrogen) to the calx of lead (minium). Thorpe points out the magnitude of the discovery that Priestley missed, and it may be said that Hales too was on the track and had he known as much as Priestley it would not have been phlogiston that kept him from becoming a Cavendish or Lavoisier. What chiefly concerns us however is the bearing of Hales' chemical work on his theories of nutrition. He concludes that "air makes a very considerable part of the substance of Vegetables," and goes on to say (p. 211) that "many of these particles of air" are "in a fixt state strongly adhering to and wrought into the substance of plants^[33]. He has some idea of the instability of complex substances and of the importance of the fact, for he says^[34] that "if all the parts of matter were only endued with a strongly attracting power, [the] whole [of] nature would then become one unactive cohering lump." This may remind us of Herbert Spencer's words: "Thus the essential characteristic of living organic matter, is that it unites this large quantity of contained motion with a degree of cohesion that permits temporary fixity of arrangement," First Principles, § 103. With regard to the way in which plants absorb and fix the "air" which he finds in their tissues, Hales is not clear; he does not in any way distinguish between respiration and assimilation. But as I have already said he definitely asserts that plants draw "sublimed and exalted food" from the air.

As regards the action of light on plants, he suggests (p. 327) that "by freely entering the expanded surfaces of leaves and flowers" light may "contribute much to the ennobling principles of vegetation." He goes on to quote Newton (Opticks, query 30): "The change of bodies into light, and of light into bodies is very conformable to the course of nature, which seems delighted with transformations." It is a problem for the antiquary to determine whether or no Swift took from Newton the idea of bottling and recapturing sunshine as practised by the philosopher of Lagado. He could hardly have got it from Hales since Gulliver's Travels was published in 1726, a year before Vegetable Staticks. Timiriazeff, in his Croonian Lecture^[35], was the first to see the connexion between photosynthesis and the Lagado research.

Nevertheless Hales is not quite consistent about the action of light; thus (p. 351) he speaks of the dull light in a closely planted wood as checking the perspiration of the lower branches so that "drawing little nourishment, they perish." This is doubtless one effect of bad illumination under the above-named conditions, but the check to photosynthesis is a more serious result. In his final remarks on vegetation (p. 375) Hales says in relation to greenhouses, "it is certainly of as great importance to the life of the plants to discharge that infected rancid air by the admission of fresh, as it is to defend them from the extream cold of the outward air." This idea of ventilating greenhouses he carried out in a plant house designed by him for the Dowager Princess of Wales, in which warm fresh air was admitted. The house in question was built in 1761 in the Princess's garden at Kew, which afterwards became what we now know as Kew Gardens. The site of Hales' greenhouse, which was only pulled down in 1861, is marked by a big Wistaria which formerly grew on the greenhouse wall. It should be recorded that Sir W. Thiselton-Dyer^[36] planned a similar arrangement independently of Hales, and found it produced a marked improvement of the well-being of the plants.

It is an illuminating fact that though Hales must have known Malpighi's theory of the function of leaves (which was broadly speaking the same as his own), he does not as far as I know refer to it. In his preface, p. ii, he regrets that Malpighi and Grew, whose anatomical knowledge he appreciated, had not "fortuned to have fallen into this statical^[37] way of inquiry." I believe he means an inquiry of an experimental nature, and I think it was because Malpighi's theory was dependent on analogy rather than on ascertained facts, that it influenced Hales so little.

There is another part of physiology on which Hales threw light. He was the first I believe to investigate the distribution of growth in developing shoots and growing leaves by marking them and measuring the distance between the marks after an interval of time. He describes (p. 330) and figures (p. 344) with his usual thoroughness the apparatus employed: this was a comb-like object, shown in Plate IX, made by fixing five pins into a handle, ¹⁄₄ inch apart from one another: the points being dipped in red-lead and oil, a young vine-shoot was marked with ten dots ¹⁄₄ inch apart. In the autumn he examined his specimen and finds that the youngest internode or "joynt" had grown most, and the basal part having been "almost hardened" when he marked, had "extended very little." In this—a tentative experiment—he made the mistake of not re-measuring his plants at short intervals of time, but it was an admirable beginning and the direct ancestor of Sachs'^[38] great research on the subject.

In his discussion on growth it is interesting to find the idea of turgescence supplying the motive force for extension. This conception he takes from Borelli^[39].

Hales sees in the nodes of plants "plinths or abutments for the dilating pith to exert its force on" (p. 335); but he acutely foresees a modern objection^[40] to the explanation of growth as regulated solely by the hydrostatic pressure in the cell. Hales says (p. 335): "but a dilating spongy substance, by equally expanding itself every way, would not produce an oblong shoot, but rather a globose one."

It is not my place to speak of Hales' work in animal physiology, nor of those researches bearing on the welfare of the human race which occupied his later years. Thus he wrote against the habit of drinking spirits, and made experiments on ventilation by which he benefited both English and French prisons, and even the House of Commons; then too he was occupied in attempts to improve the method of distilling potable water at sea, and of preserving meat and biscuit on long voyages^[41].

We are concerned with him simply as a vegetable physiologist

Plate IX

Plate 18 from Hales's Vegetable Staticks

Fig. 40. Instrument devised by Hales to make prick-
marks on a young shoot of Vine (Fig. 41); the
distribution of stretching after growth is shown in
Fig. 42. The use of a similar instrument for
marking surfaces is shown in Figs. 43 and 44

and in that character his fame is imperishable. Of the book which I have been using as my text, namely, Vegetable Staticks, Sachs says: "It was the first comprehensive work the world had seen which was devoted to the nutrition of plants and the movement of their sap....Hales had the art of making plants reveal themselves. By experiments carefully planned and cunningly carried out he forced them to betray the energies hidden in their apparently inactive bodies^[42]." These words, spoken by a great physiologist of our day, form a fitting tribute to one who is justly described as the father of physiology.

1. In 1699 Newton was made master of the Mint and appointed Whiston his Deputy in the Lucasian Professorship, an office he finally resigned in 1703 (Brewster's Life of Newton, 1831, p. 249).
2. "There, if anywhere, his dear shade must linger," Trevelyan, Life and Letters of Lord Macaulay, 1 volume edit. 1881, p. 55.
3. Black's discovery of CO₂, however, was published in 1754, seven years before Hales died, but Priestley's, Cavendish's and Lavoisier's work on O and H was later.
4. 1837, III. p. 389.
5. Vegetable Staticks, p. 346.
6. Sachs, Geschichte, p. 502. Malpighi held similar views.
7. Ibid., p. 499.
8. Quoted by Caröe, in his paper read before the Cambridge Archaeological Society on King's Hostel etc., and "Printed for the Master and Fellows of Trinity Coll." in 1909.
9. He also held the living of Farringdon in Hampshire where he occasionally resided.
10. Dict. Nat. Biog.
11. With a certain idleness Pope reduces him to plain Parson Hale, for the sake of a rhyme in the Epistle of Martha Blount, I. 198.
12. The original reads "deigned not," an obvious slip.
13. This he does by means of a network of threads ¹⁄₄ inch apart. Pfeffer, Pflanzenphysiologie, ed. I, I. p. 142, recommends the method and gives Hales as his authority.
14. Pflanzenphysiologie, 1865 (Fr. Trans. 1868), p. 254.
15. He gives it as 15.8 square inches, the only instance I have come across of his use of decimals.
16. Arbeiten, II. p. 182.
17. See Sachs' Pflanzenphys. 1865 (Fr. Trans. 1868), p. 257, where the above correction is applied to Hales' work.
18. Vegetable Staticks, p. 5.
19. Vegetable Staticks, p. 14.
20. Vegetable Staticks, p. 41.
21. Janse in Pringsheinis Jahrb. XVIII. p. 38. The later literature is given by Dixon in Progressus Rei Bot. III., 1909, p. 58.
22. Compare F. von Höhnel, Bot. Zeititng, 1879, p. 318.
23. This is also shown by experiment XC, Vegetable Staticks, p. 123.
24. The method by which Hales proposed to record the depth of the sea is a variant of this apparatus.
25. Vegetable Staticks, p. 92.
26. According to Sachs (Geschichte, p. 509) Ray employed this method.
27. Other facts show that the "gapped" branches did not behave quite normally.
28. He refers (p. 141) to what is in principle the same experiment (see Fig. 27) as due to Mr Brotherton, and published in the Abridgment of the Phil. Trans. II. p. 708.
29. He notices that the swelling of the bark is connected with the presence of buds. The only ring of bark which had no bud showed no swelling.
30. It appears that Mayow made similar experiments. Dict. Nat. Biog. s.v. Mayow.
31. History of Chemistry, 1909, I. p. 69.
32. Hales made use of a rough pneumatic trough, the invention of which is usually ascribed to Priestley (Thorpe's History of Chemistry, I. p. 79).
33. He speaks here merely of the apples used in a certain experiment, but it is clear that he applies the conclusion to other plants.
34. Vegetable Staticks, p. 313. It should be noted that Hales speaks of organic as well as inorganic substances.
35. Proc. R. Soc. LXXII., p. 30, 1903.
36. The above account of Hales' connexion with the Royal Gardens at Kew is from the Kew Bulletin, 1891, p. 289.
37. I am indebted to Sir E. Thorpe for a definition of statical. "Statical (Med.) noting the physical phenomena presented by organised bodies in contradiction to the organic or vital." (Worcester's Dictionary, 1889.)
38. Arbeiten, I.
39. Borelli, De Motu Animalium, Pt II. Ch. xiii. According to Sachs, Ges. d. Botanik, p. 582, Mariotte (1679) had suggested the same idea.
40. Nägeli, Stärkekorner, p. 279.
41. See his Philosophical Experiments, 1739.
42. Geschichte d. Botanik, p. 515 (free translation).