The plants inclosed at Arlington produced 0.44 pod to the raceme more than the plants inclosed at Ames, and the average for the six plants at Arlington and at Ames is only 0.42 pod to the raceme. Results given below for nine plants inclosed in the glass-covered cage show that the pods produced per raceme by different plants varied from 0.1 to 0.45, which is slightly less than the variation in the two cages covered with cheese-cloth.

In order to determine whether the shading of the plants in the cheesecloth-covered cages had caused the production of seed to be reduced, a cage 4 feet wide, 4 feet high, and 10 feet long, having glass sides and top, but with ends covered with cheesecloth to permit ventilation, was placed over nine plants at Ames in August, 1916. The results obtained in this experiment are presented in Table IV.Table IV.—Production of sweet-clover seed by plants protected from insect visitation during their entire flowering period at Ames, Iowa, in 1916.

The results given in Table IV show that an average of 0.31 of a pod to the raceme was obtained from 1,594 racemes and that the variation in seed production of the different plants was from 0.1 to 0.45 to the raceme. The average seed production for the nine plants is 0.11 seed to the raceme less than the average results obtained from the six plants that were covered with cheesecloth. As this difference is well within the limit of variation for individual plants, it may be stated that the shading of the plants in the cheesecloth-covered cages did not reduce the production of seed. The results of this experiment show that spontaneous self-pollination does not occur regularly, as stated by Kirchner.

FLOWERS POLLINATED ONLY BY NIGHT-FLYING INSECTS.

In order to determine the importance of night-flying insects as pollinators, two cheesecloth-covered cages 3 feet square and 3½ feet high were placed over sweet-clover plants at Arlington on July 10, 1916. The covers of the cages were removed each evening at 7:30 and replaced each morning at 4:30 o'clock. Practically all the flowers on these plants had bloomed by August 2, and the seed produced was nearly mature. The few racemes that contained opened flowers or buds were discarded. The three plants in one cage produced 723 racemes, with an average of 3.76 pods each, while the one plant in the other cage produced 227 racemes, with an average of 3.58 pods to the raceme. The four plants, therefore, produced a total of 950 racemes, with an average of 3.71 pods each. The only night-flying insect found working on sweet clover while these plants were in bloom was Diacrisia virginica Fabr.

This experiment was duplicated at Ames in August, 1916, with the result that one plant subject to visitation only by night-flying insects produced 486 racemes, with an average of 16.5 pods each.

The results obtained in these experiments show that night-flying insects were much more active in pollinating sweet clover at Ames than at Arlington. However, as the results obtained from the plants subject to visitation by day-flying insects only were practically the same as those obtained from plants which were subject to insect visitation at all times, it is concluded that night-flying insects were not a factor in the pollination of sweet clover at Arlington or at Ames in 1916.

FLOWERS POLLINATED ONLY BY DAY-FLYING INSECTS.

A cheesecloth-covered cage, 3 feet square and 3½ feet high, was placed on July 7, 1916, over two sweet-clover plants at Arlington, before any of the flowers opened. As the cover of this cage was removed at 7.30 a. m. and replaced at 4.30 p. m. each day during the experiment, the plants were subject to visitation by day-flying insects only. As soon as all of the flowers on most of the racemes had bloomed, and before any mature pods shattered, the racemes were removed from the plants and the pods produced by each raceme counted. The two plants produced a total of 544 racemes, with an average of 20.9 pods each.

This experiment was also conducted at Ames. One plant was protected from insect visitation at night in August, 1916, with the result that it produced 418 racemes, with an average of 41.11 pods each.

PLANTS PROTECTED FROM ALL INSECTS THAT COULD NOT PASS THROUGH A WIRE SCREEN HAVING 14 MESHES TO THE LINEAR INCH.

It is well known that many small insects, and especially those belonging to the family SyrphidÆ and to the genus Halictus, frequent sweet-clover flowers, but no records have been noted that show how important these insects are as pollinators of this plant. In order to obtain data on this subject a cage 12 feet square and 6½ feet high, made of wire screen having 14 meshes to the linear inch, was placed over a few plants at Ames, in July, 1916, before they began to bloom. The base of the cage was buried several inches in the soil, so that no insects could get into it. As these plants were growing in a field where there was a sufficient supply of moisture at all times, they made a growth of 5 to 6 feet. For this reason all the racemes were collected from only a portion of one of the plants instead of from the entire plant, as was done with the smaller ones inclosed in the cheesecloth-covered cages. The branches selected contained 224 racemes, with an average of 24.53 pods each. Many insects that were able to pass through the wire netting were observed working on the flowers of the inclosed plants.

A check plant, subject to visitation by all insects and growing within a few yards of the cage, contained 264 racemes, with an average of 28.23 pods each.

This experiment shows that small insects are efficient pollinators of sweet clover and that the plant to which all insects had access produced an average of only 3.7 pods to the raceme more than the one inclosed in the cage. As these plants were growing close to a strip of timber and some distance from a field of sweet clover, it is probable that more small insects worked on the flowers than would have been the case if the cage had been located in the center of a field of sweet clover. Though these results show that small insects are able to pollinate sweet-clover flowers freely, it is very doubtful whether insects of this kind would be numerous enough to pollinate sufficient flowers in a large field of sweet clover for profitable seed production. The honeybee is the most efficient pollinator of this plant, and it is believed that in many sections it is responsible for the pollination of more than half of the flowers.

SUMMARY OF INSECT-POLLINATION STUDIES.

The data secured in the different experiments where sweet-clover flowers were subject to insect visitation at one time or another are presented in detail in Table V.Table V.—Summary of the insect pollination studies conducted at Arlington, Va., and Ames, Iowa, in 1916.

The results in Table V show that an average of 0.37 pod to the raceme was obtained from the plants protected from visitation by all insects during the flowering period. As the racemes of Melilotus alba will average approximately 50 flowers each, less than 1 per cent of them set seed without being pollinated by insects. The results obtained in the cages in which only night-flying insects had access to the flowers show that these insects pollinate sweet clover to a slight extent, but that the number of pods produced by them is so few that it may be assumed that these flowers would have been pollinated by day-flying insects. This assumption is borne out by the results obtained in the cages where only day-flying insects had access to the flowers, as the results obtained in these cages at Arlington and Ames, respectively, are approximately the same as those obtained on the plants subject to insect visitation at all times. It will be noted that the yield of seed on the plants visited by insects at Ames is much higher than that of the plants subjected to insect visits during the same period at Arlington. This difference in seed yield may be attributed to the fact that isolated plants were used in the experiments at Ames, and at Arlington the experiments were conducted with plants growing under field conditions.

RELATION OF THE POSITION OF THE FLOWERS ON MELILOTUS ALBA PLANTS TO SEED PRODUCTION.

Observations of sweet-clover plants grown under cultivation, and especially when the stands were thick, showed that the flowers of the racemes on the upper and exposed branches produced a larger percentage of seed than those on the lower branches which were less exposed. It is thought by some that the failure of the flowers on the lower racemes to be fertilized is due to shading; but the results obtained in the cheesecloth and glass covered cages do not warrant this belief, as it is doubtful whether the shading of the flowers on the lower racemes is more than that caused by the cheesecloth. It is probably the lack of pollination that causes this decrease in seed production on the lower branches of plants growing close together, as a vast number of flowers open each day on portions of the plants which are exposed directly to visitation by insects and are therefore more accessible to them.

In order to obtain information upon the number of flowers that produce seed on the upper and lower portions, respectively, of sweet-clover plants when grown under field conditions and where the stand contained four to five plants to the square foot, a number of racemes were labeled on different portions of the plants at Ames in 1915 and 1916. When the pods were partly mature, records were made of the number of flowers that produced pods. The results obtained are given in Table VI.Table VI.—Relation of the position of sweet-clover flowers on the plants to seed production, at Ames, Iowa, in 1915 and 1916.

The flowers on the upper racemes of the plants produced 31.2 per cent more pods than those on the lower racemes in 1915. and 11.4 per cent more in 1916. These results prove that insects more frequently visit the flowers that are directly exposed and are therefore more accessible.

INFLUENCE OF THE WEATHER AT BLOSSOMING TIME UPON SEED PRODUCTION.

The seed production of sweet clover is seldom satisfactory when rainy or muggy weather prevails during the flowering period. In order to obtain data as to the relation existing between the visits of insects and the prevailing weather conditions, a record of insect visits and of the number of flowers that opened each day was kept for a period of nine days at Ames in August, 1915.

In this experiment the racemes were marked early each morning just above the last flowers which had opened the previous day, and early the following morning the number of flowers which had opened the previous day was noted. The number of flowers that were pollinated was determined by the number of pods that formed. Table VII gives in detail the results obtained.Table VII.—Influence of the weather at blossoming time upon the yield of sweet -clover seed, at Ames. Iowa, in 1915.

The data given in Table VII show that the percentage of effective pollination is much higher in clear weather, when insects are active, than in cloudy or rainy weather, when but few insects visit the flowers.

INSECT POLLINATORS OF SWEET CLOVER.

On account of the ease with which the heavy flow of nectar of sweet-clover flowers may be obtained many insects visit the flowers, thereby pollinating them. While the useful insect visitors of flowers of red clover are limited to a few species of Hymenoptera, those pollinating sweet-clover blossoms are many and belong to such orders as Coleoptera, Lepidoptera, and Diptera, as well as to the Hymenoptera. However, in the United States the honeybee is the most important pollinator of sweet clover. In many parts of the country the different species of Halictus, commonly known as sweat bees, rank next in importance. The margined soldier beetles (Chauliognathus marginatus Fabr.) were very active pollinators at Arlington, Va., in the latter part of June and first part of July, 1916, but the woolly bear (Diacrisia virginica Fabr.) was the only night-flying insect found working on sweet clover at Arlington.

Insects belonging to the genera Halictus, Syritta, and Paragus were very active pollinators at Ames, Iowa, in 1916, and ranked next in importance to the honeybee. In fact, the results obtained in the cage where the plants were protected from visitation by insects that could not pass through a screen having 14 meshes to the linear inch showed that these small insects were able under the conditions of that experiment to pollinate practically as many flowers as larger insects.

The insects listed below were collected while visiting Melilotus alba and M. officinalis flowers in 1916.

AT ARLINGTON, VA.

AT AMES, IOWA.

EFFECT OF MOISTURE UPON THE PRODUCTION OF MELILOTUS ALBA SEED.

Careful inspection of a number of sweet-clover fields in Iowa and Illinois in the autumn of 1916 indicated that many plants were unable to obtain sufficient moisture for the proper development of their flowers. Examination of flowers that aborted shortly after reaching their mature size showed that the anther sacs had not burst, even though the pollen grains were mature. Apparently for the same reason many immature pods aborted. The precipitation for July, 1916, in Livingston County, Ill., where the sweet-clover seed crop suffered materially for lack of moisture, was 3.2 inches less than normal, while the temperature was 4.5° F. above normal. In August the precipitation was 0.96 of an inch below normal and the temperature 4.2° F. above normal. At Ames, Iowa, the precipitation was 3.54 inches below normal and the temperature 5.4° F. above normal in July. Both the precipitation and temperature were about normal at Ames in August, but most of the precipitation fell before the experiments were commenced.

In north-central Illinois the seed production of sweet clover was very irregular. Some fields produced an abundance of seed, while a large percentage of the pods on the plants in other fields near by, where the thickness of the stand, size of the plants, and conditions in general were approximately the same, aborted. It was evident that all stands producing a good seed crop were growing on well-drained soil and that those which were not yielding satisfactorily were on poorly drained land. It is well known that sweet clover will produce deep taproots only when the plants are growing in well-drained soil and that a much-branched surface root system will be formed on poorly drained land, and especially when there is an excess of moisture or a high water table during the first season's growth. During this droughty period in 1916 the upper layer of soil became so depleted of moisture that the plants with surface root systems were unable to obtain sufficient water to mature their seed. On the other hand, the lack of precipitation and the high temperatures did not affect the moisture content of the subsoil sufficiently to interfere with the normal seed production of deep-rooted plants. According to Lutts (22, p. 47) this same condition was found to be true in Ohio in 1916.

As a rule, under droughty conditions the second crop of sweet clover will produce a higher yield of seed than the first crop, as the second growth of the plants is seldom more than half as much as the first, thereby requiring less moisture. However, if showery hot weather prevails when the first crop is cut, the end of each stub is very apt to become infected, usually with a species of Fusarium, which kills all the cortex as far back as the upper bud or young shoot and that part of it on the opposite side of this bud to the bud below. If the second bud from the top of a stub is not directly opposite the upper one the decay may extend nearly to the ground. (Pl. IV.) The destruction of half to two-thirds of the cortex from 2 to 4 inches below the upper bud materially reduces the quantity of water that can be conveyed to the branch above the base of the dead area. Plants thus infected obtain sufficient moisture for seed production only under the most favorable conditions. When the first crop is cut during warm dry weather, and especially when the first crop has not been permitted to make more than a 30 to 32 inch growth, the stubble seldom decays, and in no instance have the plants been observed to decay as far back as the upper buds.

An experiment was conducted at Ames in the latter part of August and first part of September, 1916, to determine the effect of watering plants that were aborting a large percentage of their flowers and immature pods. For this purpose several volunteer plants growing in a meadow were selected. A hole 12 inches square and 14 inches deep was dug 8 inches from the crown of one plant, and each evening during the experiment 2 gallons of water were poured into the hole. The top of the hole was kept covered, so as to check evaporation from it as much as possible. Another plant of the same size and growing about 15 yards from the watered plant served as a check. On both plants many of the flowers and immature buds were aborting at the beginning of the experiment. The soil in this field was so depleted of moisture that the leaves of the plants wilted during the hottest part of the days preceding the experiment. The foliage on the check plant wilted each day for the first five days of the experiment. On the sixth day 0.96 of an inch of rain fell and four days later 0.23 of an inch more. The dropping of the flowers was temporarily checked by these precipitations, but owing to the dry, compact condition of the soil the rain was not sufficient to check entirely the fall of flowers and immature pods. At the beginning of the experiment the racemes on both plants were divided into three classes, according to the development of the flowers, and labeled. They were collected and the seeds counted as soon as the pods at the bases of the racemes commenced to turn brown. Table VIII presents the results obtained.Table VIII.—Effect of water upon the seed production of sweet clover when growing under droughty conditions at Ames, Iowa, in 1916.

The effect of the water was noticeable soon after the first application, as the leaves and flowers on this plant became turgid and the anther sacs burst at the proper stage of their development. Very few flowers fell after the second day. The water decidedly checked the aborting of immature pods, as is shown by the results obtained on the racemes which were labeled after the pods had formed. The racemes which contained pods 3 to 6 days old when labeled matured 9.95 pods to the raceme more than those which contained older pods at the beginning of the experiment, but this was expected, as most of the aborting which caused this difference had taken place before the racemes were labeled. As very few pods aborted before they were 3 to 6 days old, the difference of 9.95 pods to the raceme in favor of the ones labeled when the flowers at their bases were just ready to open was largely due to the dropping of the flowers on the older racemes before the experiment was begun.

It will be seen that the production of mature pods on the plant not watered was much greater on the racemes that were labeled before the flowers opened than on the older racemes. This difference is undoubtedly due to the precipitation which fell on the sixth and tenth days of the experiment. It is believed that the yield of 15.23 pods to the raceme on the ones labeled when the pods were 9 to 12 days old is representative of the production of pods per raceme previous to the precipitation and that the other racemes on this plant would have yielded proportionately if conditions had remained the same.

In the early spring of 1916, Melilotus alba was planted in several large pots in the greenhouse of the Department of Agriculture at Washington, D. C. These pots were placed outside the greenhouse in the late spring, where they remained until the following January, when they were taken into the greenhouse. The plants grew rapidly and began to flower during the latter part of April, 1917. At this time two pots were placed in a large cage made of screen having 20 meshes to the linear inch. One pot was submerged in a tub of water, so that the soil was saturated at all times, while the plant in the other pot was given only sufficient water to keep it from wilting. The pods on a few racemes were self-pollinated and the results obtained are given in Table IX.Table IX.—Effect of moisture on the seed production of Melilotus alba at Washington, D. C, in 1917.

The results of this experiment compare favorably with those obtained under field conditions at Ames in 1916.