THE INHERITANCE OF FLOWER COLOUR IN PISUM

T H E I N H E R I T A N C E O F F L O W E R C O L O U R I N P I S U M

BY H A N S T E D I N S\'AI.fiF

1. INTRODUCTION.

Y first pea-crosses were obtained the same year I was entrusted M with the management of the breeding of peas and vetches at the Swedish Seed Association at Svalof, that is, in 1892. They were carried out with hardly any special purpose, more for my own enjoy- ment, even if not entirely without a slight hope that they might give some practical results. The numerous pea-crosses made later on in that decennium included several between different pure strains with white flowers and such with rose-coloured flowers (standard more or less reddish to white, wings light or dark rose)'. These crosses gave without exception a F, with purple flowers, and F, yielded purple, rose, and white flowers. The numerical relations were not determined, but it was noted, as is to be seen of my annotations from that time, that in an average always the purple-coloured flowers were in a decided majority in proportion to each of the other two. The different pure strains with rose flowers were also crossed with each other, but however they were combined they never gave purple or white flowered individuals, neither in F, nor in the following generations.

In 1901 one of the rediscoverers of the ))Mendelian laws)), Pro- fessor E. TSCHERMAK from Vienna, visited Svalof. On that occasion I gave him 10 different pure strains of Pisum arueme and 9 different

Already at that time a very large number of pure strains of peas were grown at Svaliif includi,ng such with white flowers as well as such with purple flowers (standard more or less light purple, wings dark purple). All these pure strains were selected by the pedigreemethod from native or foreign mixed sorts or found as intermixtures in other kinds of corn etc. I also had five different pure strains with rose flowers (See HANS TEDIN och HUGO WITT: Botanisk-kemisk undersokning av 42 n8stan uteslutande nya lirtformer, uppdragna vid Sveriges Utsf. pi3 Svaliif. - Sveriges Utsf:s Tidskrift 1899, pages 121-160). The 42 strains descri- bed in this paper were but a little collection of those selected which were cultivated or stored in the collection of the Seed-Ass.

THE INHERITANCE O F FLOWER COLOUR I N PISC'M 69

F, crosses of which four represented crosses between white and rose flowered peas (TSCHERMAK 1902). For these he (1904 a and b) pointed out segregation in F, in the ratio 9 : 3 : 4 - facts which now are well known, and which afterwards have been pointed out by several workers. Further examples will be given in this paper. On the presence and absence hypothesis this fact is interpreted thus: the white-flohering pea has a cryptomeric factor which alone has no visible effect, but which combined with the rose factor gives purple flowers. If the rose factor is missing the flowers will be white whether the other factor is present or not. As far as I can find these two flower colour factors in Pisum are the only ones that have been pointed out in previous literature. WHITE (1917 a ) thus mentions only two: A (,Salmon pink or rose flower colourn) and B (xpurpling factor; gives with A purple flowers,). LOCK (1908) names the same factors C and P, TSCHERMAK (1912) in conformity with WHITE A and B andKAJANUs (1919) R and G.

2. MATERIAL AND METHODS.

Already in 1898 I received from the Agricultural College of Ultuna, Sweden, a pea called PLjusrod-blommig GrHartn, i. e. a light red flowered grey pea (ngrey pea, is the Swedish name for Pisum aruense), a sort of pea with flowers as pale as the rose-coloured, and, accordingly, with white or almost white standard, but with wings not rose-coloured but light purple (See plate I). This pea will be named in he following by its stock-book number, 01001. Several other lines mentioned in this paper will receive similar numbers. When crossing this 01001 with natural whites, i. e. white forms not obtained by artificial crosses, I received without exception a purple flowered F1, just as when crossing white and rose. But in F, not only purple, light purple and white flowers appeared, but also rose and further a fifth colour, namely violet (see plate I) . Phenotypically this violet can be said to stand in the same relation to purple as light purple to rose.

A closer examination of the colour inheritance in Pisum could not for several reasons be started until 1911. Although the working out of the different modes of inheritance at first seemed to be very simple, serious difficulties were met with in the course of the investigations.

Moreover, only limited time and work could be given to these experiments, the breeding for practical use requiring most of the time and labour.

Difficulties are often met with in the marking of the different

70 HANS TEDIN

flower colours in the field due to the great modifiability of the plants. The wings of the light purple flowers change sometimes their colour to nearly white with only a pale shade of light purple in the edges. They get then easily confounded with the whites. 'l'he purple and rose coloured flowers are modified in the same way. W H ~ T E (1. c.) says: ,,weather conditions, especially prolonged damp or rainy weather, often wash out or suppress the development of the purple flower colour, so that it resembles the pink: and pinks are modified by the same causes to whiten. I have never observed such a marked modifi- cation of purple and rose, however, nor have I any decided opinion as to the cause. Last year, for instance, the first developed flowers of 01001 (first part of July) were almost white, while the following ones (latter part of July to late in the autumn) developed the light purple colour quite typically, although the weather was mostly cloudy with hardly any sunshine and at times with heavy rains. When growing older the purple and rose coloured flowers get a t bluish tinge. They are now easily confounded with the violet and light purple flowers. Hence the marking of flower colour should always be done on flowers just opened,

Very much damage and very great difficulties have been caused by thrips, the small injurious insects belonging to Thysanoptera. Its attacks on the buds often prevent them from opening. In this way all the F, and F, material in 1914, 1916, 1917 and 1918 was destroyed. Some of the F, and F, families are lost or more or less damaged every year, due to the attacks of these insects. No reliable figures can be obtained from such material, and these families have to be discarded. Nevertheless individuals from certain of these F, progenies have been selected and sown to F, in some cases where the flower colour could be determined with certainty. This explains, for instance, the circumstance that the number of purples sown in 1912 (tab. 3-9) exceeds the number of purples obtained in F, 'in 1911 (tab. 1).

The sowing was done by hand in four rows with 20 cm. between the rows and 10 cm. between each grain in the row; twigs were used as support. Although the plants stood so closely and were entwined in*eack other the determining of the number of plants in the different colour classes segregated could be done without greater difficulties, if only two qualified persons were helping each other; for one person it is hardly possible.

The difficulties stated in the above make it evident, that it is almost impossible to avoid errors in the marking. Besides the confu-

T H E INHERITANCE OF F L O W E R COLOUR IN PISUM 7 1

sion of flower colours it also happens that a stout basal branch, which can be followed to the surface of soil, is mistaken for an individual plant. Because of my absence at certain times it has not always been possible to control the work as strictly as could be wished. Therefore, there is no doubt that just in the errors made in the marking we will find the most important cause to the fact that the numerical relations (especially of the descendants of the particular plants) not always agree with the theoretical ratio as well as could be wished. The number of the individuals ha's often been small, and this fact explains some cases of nonconformity. Last year the marking was made especially difficult by a very irregular and uneven development of the plants.

It is evident that the marking of the plants would be both easier and more accurate, if they were sown more sparsely, in two or still better in only one row with greater distance between the plants in the row. But both space and expenses are saved by the way of sowing used in my cultures, and the development of the plants is further kept within more normal limits. If some care is taken, no difficulty is met with in separating the plants without tearing them when harvesting.

The investigations are not yet completed. (This year a great F, and various F, crosses will be sown). Nevertheless I have for several reasons thought i t better not to put off the account of the results already obtained. I hope, however, in course of time to have occasion to come back to these questions in connection with an account of an investiga- tion already started on the genetic relations between the flower colour and leaf axils and certain characteristics of the seeds.

This paper deals exclusively with the flower colours.

3. THE RESULTS OF THE INVESTIGATIONS.

A s far as can be observed from the investigations already carried out the mode of inheritance of the flower 'colours in Pisam is very simple, as mentioned before, although different from the one already knQwn. Crosses between 01001 (homozygous light purple) and common whites give without exception purple flowered F1, and F, segre- gates in purple, violet, rose, light purple and white in the ratio 27 : 9 : 9 : 3 : 16 (table 1). From what is said in the above and consi- dering'the segregation of the progenies in F, (fables 2-16) as well as the results of the back-crosses and other crosses with whites from

72 HANS TEDIN

F, (see pag. 76-77 and tables 17-20) it is at once observed, that purple colour is conditioned not only by two factors but by three. The pre- sumtive factor for rose adopted from previous investigations (WHITE’S and TSCHERMAK’S factor A, meiitioned before) is n o t a simple factor but composed of two factors. One of these, which I call A, gives light purple, and this one is also the real fundamental factor for the flower colour in Pisum aruense. A second factor, my B factor, gives rose together with A. Then there iS an additional third factor C (WwrE’s and TSCHERMAK’S purpling factor B) which together with A gives violet, while all the three together give purple. B and C are, therefore, both alone or coexisting without effect (cryptomeric or masked) as soon as A is absent.

Thus the different flower colours in pens show the Following constitution in the homozygous states according to my results so far obtained.

Light purple: AAbbcc. Rose: AABBcc. Violet: AAbbCC. Purple: AABBCC. White:

No difference between homo- and heterozygotes of the respective flower colours has been observed.

The whites crossed with the homozygous purple (AABBCC) by MENDEL and other workers must have had the constitution aaBBCC as a segregation in F, in purple and white in the ratio 3 : 1 invariably took place. And the whites used hy various workers when crossing with the homozygous rose (AABBcc) must have been constituted in the same manner as the segregation in F, resulted in purple, rose and white in the ratio 9 : 3 : 4, thus necessitating a F, constituted AaBBCc. These observations favour the belief that the common white flowered varieties of the peas usually if not always (most of them have not yet been investigated in this respect) contain both B and C in homozygous state. This circumstance would get quite a natural explanation if we, following the opinion of LOCK (1. c.), suppose that the whites have originated from purple flowered plants by the ,loss)) of the colour factor A, a supposition quite hypothetical a t present.

It is of very great interest to find that the hilum of the violet flowered individuals have an abnormal structure, and these plants show also a very poor development of the seeds. A normally developed

aaBBCC, aabbCC, axBBcc and aabbcc.

T H E INHERITANCE O F F L O W E R COLOUR IN PISUM 73

hilum has an oval shape. Outmost it consists of a layer of thick walled palisade like cells (Hpl) sunk in a shallow depression of the external layer of the testa, (see fig. 1 which shows a cross section of n normal hilum), which also consists of thick walled pahsade cells (Tpl).

Fig. 1. Cross section of normal hilum. - TpZ: the outermost layer of the testa composed of thickwalled palisade cells. - Hpl: layer of palisade cells belonging to hilum. - S: the slit in the

middle of the hilutn. - Tr: tracheal tissue.

Both the palisade layers decrease in size towards the slit (S) in the middle of the hilum, and finally disappear altogether. An eggshaped or elipti- cal radially disposed tissue of tracheal elements (Tr) lies directly under the slit (see also KAJANUS 1. c.).

In the seed of the violet floweredl plants, however, hilum appears outwardly as a slitlike depression in the testa only, while the above mentioned layer con- sisting of palisade cells is lacking altogether (see fig. 2). The fact that the tracheal tissue

d /

is is Of

entirely missing Fig. 2. Cross section of hilum in violet flowered individuals. - TpZ and S same as in fig. 1. Hilum is devoid

greater con- of both palisade cells and tracheal tissue as seen in the flg. sequence. The conduc- tion of nourishment to the seeds became difficult because of the absence of the tracheal tissue, and their own development became retarded. The power of germination does not seem to be impaired even if the plantlets are somewhat weaker than those from normal seeds. On account of the

74 HANS TEDIN

poor development of the seeds on the whole it is not surprising to find a relatively large number of seeds incapable of germinating. I have not been able to find any difference with regard to these pecu- liarities between violet flowered forms in the homozygous and in the heterozygous states, nor between the seeds of the latter combination of violet coloured individuals giving rise to plants with violet or other colours (light purple or white).

The abnormal structure of the hilum is, therefore, directly connec- ted with the factorcombination AC without B. I am not able to give any explanation as to the exact n'ature of this connection, nor can I say anything about other characters possibly involved. Investigations are now being planned to settle these qnestions.

4. THE TABLES.

The results of the investigations are tabulated in tables 1-20. They contain all the detail numbers, i. e. the numerical segregation of the different F, and F, progenies. For the sake of control the field numbers of the different families are also put out (in col. 2 ) . A comparison between the observed and the theoretical segregation numbers of a rather large number of progenies from different individuals, and especially of those showing the most diverging numerical relations, has also been made, although not put out in the tables. However, the correspondence between the observed and the theoretical numbers have as a rule been satisfactory even in these. There are, nevertheless, exceptions where the deviations are too large.

Only the comparison between the theoretical numbers and the segregation numbers summed up from all of the progenies belonging together has been tabulated.

Table 1. Shows segregation in F, of three different crosses between 01001 (AAbbcc) and three different whites, two of which are without names, and the third called by its stock book number 0154. All three gave purple flowers in F,. By self-fertilizing F, a F, segregation in purple, violet, rose, light purple, and white took place in the ratio 27 : 9 : 9 : 3 : 16, as shown in the table. Consequently, the white parents must have had the constitution aaBBCC. The summed up observed and theoretical numbers agree very well. In all three crosses, however, the number of the purples is somewhat too small, while the number of whites is somewhat too large. When the numbers from

T H E I N H E R I T A N C E OF F L O W E R COLOUR IN PISUM 75

all three crosses are added (see bottom of the table) the deviation of the whites is seen to be almost three times and the deviation of the purples even more’than three times the standard error. Thus the deviation is really somewhat greater than is generally allowed.

Table 2. In this table are found (col. 2 ) all the theoretically con- ceivable factorial combinations in F, of all the flower colours except the whites, and also (in col. 3-7) the theoretical ratio of the segregation of their progenies in F,. Col. 9-14 show the numerical classi- fication in the respective factorial combinations of the F, individuals sown both theoretically (col. 10) and observed (col. 9 and 11). In col. 8. lastly, refer.ence is given to the tables (3-16) in which the numbers obtained in F, from the segregating F, individuals are listed. The observed and the theoretical numbers relating to the numerical sorting of the F, individuals in the resp. factorial combinations agree parti- cularly well. In most cases the deviation is smaller than the standard error (col. 14).

The F, individuals with rose (63 ind.) and light purple coloured flowers (55 ind.) sown in 1912 and found a t the bottom of the table do not, however, arise from the crosses I and I1 in table 1, as in the case of the purple and violet flowered F, individuals, but owe their origin from a cross 01001 X white which yielded rose flowered F, and a F, segregation of rose, light purple, and white in the ratio 9 : 3 : 4 (table 1 7 ) . Thus the constitution of F, was AaBbcc, and the white parent (from F, arising from a cross, 1907, identical with cross I in table 1) ’was constituted aaBBcc.

The fact that the rose and light purple flowered F, individuals sown to F, did not originate from the same cross as the F, individuals with purple and violet coloured flowers sown to F,, nor from a nearly like cross, may seem to be an impropriety tending to lessen the value of the investigatiw. However, this impropriety cannot be of any importance in this case. It must be assumed that the nine rose flowered F, individuals originating from AaBbcc-F,-rose are consfiuted factorially in exactly the same way as the nine rose flowered F, individuals from the crosses I and I1 and from AaBbCc-F,-purple if the explanation of the inheritance of the flower colours given in the above is correct. And that this is so is evident from the fact that their progenies (tables 13-15) have showed a segregation exactly the saine as the latter nine from crosses I and I1 theoretically calculated. And what in this case is true of the rose flowered F, individuals must also be

76 HANS TEDIN

true of the F2 individuals with light purple coloured flowers so\vn to F, (1912).

Tubles 3-16 show segregation in F, of the F, individuals sown and can, therefore, be said to be a verification of table 2. As is evident from the tables there is generally a good - in most cases a very good - correspondence between the theoretical and observed numbers in the F, generation. Table 12 and .16, however, show exception. Table 12 shows the segregation 3 violet: 1 white in the progenies of F, individuals with violet flowers and constituted AabbCC. Table 16 gives the segregation 3 light purple: 1 white in progenies of F, individuals with light purple flowers and constituted Aabbcc. In both cases the whites are in excess, and the deviation is greater than three times the standard error. Not taking into consideration those two deviations the whites are often found to be somewhat in excess. I dare not say that the whites on the whole are more branched than those otherwise! coloured, but last summer it was found that the person in charge of the marking very often overestimated the numbef of whites by coun- ting large branches as complete plants. I can give no other exp1;in;i- tion of the excess of the whites and of other deviations than that of assuming the play of incidental causes.

Table below, and tables 17-20. Back-crosses and other crosses with whites from F,.

The factorial constitution of the 16 white flowered individuals segregated in F, of crosses I, 11, an,d 111 in table 1, and the theoretica1

6 white flowered FS indi, viduals of AaBbCc-F1

Number ractorial consti, tution. Q

aaBbCc aaBbCC aaBBCc nabbCc aaBbcc aaBBCC aabbCC aaBBcc aabbcc

01001 homozygous light purple.

Theoretical results in F1

Purple Violet Rose Light purple

THE INHERITANCE OF FLOWER COLOUH I N PISL'M 77

results in F, when crossing them with 01001 are to be seen in the table on the preceding page.

Thus, nine whites differently constituted and nine different possibilities in F,. I have carried out 46 such crosses in all, and I have found all the nine possibilities realized. However, in some cases the number of individuals in F, has been rather small. It is my intention to carry out such crosses in a great extent for control.

Two of the above mentioned 46 crosses have been followed up in F,. One of them is the one mentioned in table 17, whose white parent were constituted aaBBcc. Another white (from F, of cross I, table 1) yielded light purple F,, and light purple and white in the ratio 3 : 1 in F,, and were, therefore, constituted aabbcc (table 18).

A third white (from cross I, table 1) yielded in both cases, when crossed with two homozygous purple (0401 and 0351), purple flowered F,, and in F, purple, rose, and white in the ratio 9 : 3 : 4 (table 19). Thus, F, was AaBBCc and the white parent aaRBcc, just as in the cross in table 17.

Finally, a fourth white (also from cross I, table 1) yielded when crossed with 01412 (homozygous rose AABBcc) purple F,, and in F, the same segregation took place as in the case of the last mentioned cross (purple, rose, and white in the ratio 9 : 3 : 4) . Consequently, this white had the usual constitution of ordinary white or aaBBCC (table 20).

Lastly, I have also crossed 01001 with the two above mentioned homozygous purple (0401 and 0351). In both cases F, segregated in purple, violet, rose, and light purple, as was to be expected. Althoiigh the numerical relaltions could not be ascertained with sufficient exact- ness on account of the attacks of thrips, the segregation of these two crosses in F, speak also strongly in favour of the correctness of the results otherwise arrived at in my study of the inheritance of the flower coloiirs in Pisum.

78 HANS TEDIN

I ))

n n

TABLE 1. Cross I: 01001, AAbbcc, light purple 9X aaBBCC, white (nameless) d.

n 11: )) )) )) x )) ))

)) 111: lo 1) )) x u )) 0154. FI in all three cases purple AaBbCc.

1911 337 )) 343 )) 349 u 352

S e g r e g a t i o n i n Fs Cross I Year 1

numbei Total Light

purple White Purple Violet Rose

4 6 6 7 8 9

29 28 39 39 135 64 64

-

-

- - - 64 11 68 15 35 8 34 26 24 18 21 32 356 64 64

-

- - -

42 7 8

7 9 13 8 37 16 17,541 + 1,541

f 2485

15 1 17 8 6 3 13 12 8 6 4 10

0,646

7 9 16 13 45 27

- 5,087

- 21,838

f 2,720 2,083

23 5 32 3 20 2 8 9 9 3 4 13 131 27 23,550

- 3,410 f 1,an

2,000

12 3 6

6 3 3 11 23 9 10,804 + 1,904

f 1,915 0,994

13 3 9

4 2 4 2 2 4 7 3

-

9 1 6 5 24 9 1 1,378 + 2,878 1,242

f 1,916

9 1 10 3 5 1 4 2 3 2 5 1

- 3 1 2 6 3

- 0,186 2,844

f 1,164 0,134

4 1

1 -

- - 5 1 2 3 1 5

531 532 533 534 535 536 537 538 539 540 541 542

rota1 ..................... 3atio ..................... 'ound ..................... Ieviation, D ............ itandard error, M, )/M, ....................

53 9 9,628

4- 0,528 f 1,179

0,448

3 1 -

46 9 8,270

- 0,780 f 1,179

0,618

9

1 -

23 3 4,185 + 1,135

f 0,717 1,583

- 2 -

103 16 18,617 + 2,617

f 1,469 1,713

18 1 1

I11 ))

)3

1919 u n

378 379 380

THE INHERITANCE OF FLOWER COLOUR IN PISUM 79

- 3

381 382 383 384 385 386 387 388 389 390 39 1 392 393 394 395 396 397 398 399 400 401 402 403 404 405 406 407

-

rota1 ..................... Ratio ..................... Found ..................... Deviation, D ............ Standard error, M, D/M, .....................

Sum total ............... Ratio ..................... Found ..................... Deviation, D ............ Standard error, M, DIM, .....................

4

4 4

11 4

13 9 1 8

10 6 5 9 8 6

11 11 8 2

17 11 2 2 1

11 5 7

11 218 27 24,826

- 2,174

f 1,333

1,6SI

45 131 218 394 27 23,847

- 3,053 f 0,974

3,135

5

2 1 1 - - 2 1 4 2 2

9 2 1 6 2

-

- - 6 3 1 2 1 3 5 3

11 74 9 8,427

f 0,838

23 53 74

150 9

- 0,573 0,611

9,117 $- 0,117

f 0,686

0,171

6

- 2 3 1 4 1 1 2 1 2

6 5 2 5 4 8

7 5 1 3 3 3 1 2 1

-

-

83 9 9,451 + O , m

f 0,838 0,482

24 46 83

153 9 9,299

f. 0,686

0,436

+ 0.289

7

22 3 2,506

& 0,571

0,867

- 0,495

6 23 22 51 3 3,100

f. 0,417 0,240

+ 0,100

8

1 2 3 3 5 5 4 3 6 5

21 4 4 4 10 4 1

13 5 2 7 2 7 4 9

11

-

165 16 18,790

$. 2,790

2,387 f 1,169

37 103 165 305 16 18,537 + 29537

f. 0,654

2,871

- 9 - 7 9

18 8

24 17 8

17 20 18 5

47 21 13 27 28 20 3

45 25 6

15 7

23 15 22 35

562 64 64 - - -

135 356 562

1053 64

- - - - -

80 HANS TEDIN

TABLE 2. Factorial constitution of Fdndidduals and theoretical segregation of their progeny.

~ ~~

Flower colour and factorial constitution 3f Fs-individ.

2

Purple: AaBbCc AABbCc AaBbCC AaBBCc AABbCC AABBCc AaBBCC AABBCC

Purple : AaBbCc AABbCc AaBbCC AaBBCc AABbCC AABBCc AaBBCC AABBCC

Violet: AabbCc AAbbCc AabbCC AAbbCC

Violet: AabbCc AAbbCc AabbCC AAbbCC

Rose: AaBhcc AABhcc AaBl3cc AABBcc

Light Purple: Aabbcc AAbbcc

T h e o re t i C P 1 r a t i o

3 4

II-

- 3 11 4 8 5 9 6 5

- 1 8 6 9 5

- 2 47

3 31 4 8 5 15 6 11 7 5 8 10 9 9

- 5 94

10 13 11 6 - 0 - 3

22

10 14 11 8

- 5 37

13 33 14 9 15 12 - 9

63

16 39 - 16

55

-

-

-

12 10 -

-

-

- .-. 5 g ::

4 L

10 - - 8 4 4 4 2 2 2 1

27

8 4 4 4 2 2 2 1

27

4 2 2 1 9

4 2 2 1 9

4 2 2 1 9 2 1 3

-

-

-

-

-

-

11 11 12 I 13 1 14

6,319 - 1,681 f 1,798 0,93!

4,596 +0,596 f l , 3 9 9 0,421

2,872 -1,izs f1,ws 0,801 0,575 -1,425 f1 ,031

2,872 +0,872 f 1,091 0,841

5,170 f 1,170 f 1,399 0,831

3,447.+1,447 f l , O 3 l 1,40:

- 1,149 +O,l49 f0 ,744 0,201

7,000

8,904 +O,904 f 1,272 0,71

2,298 - 1,702 f 0,989 1,72

4,309 +0,3us fO,s89 0,31:

3,160 - 0,840 f 0,988 0,841

1,498 -0,564 fO,729 0,771

2,872 +0,872 f0 ,728 1,191

2,585 +O,585 f O , m (),so: - 1,436 +O,436 f0 ,526 0,821

7,000

5,318 +1,318 fO.953 1,SS:

2,455 +0,455 f0 ,798 0,571

0,000 - 2,000 f 0,798 2,501 - 1,aw +0,227 f0 ,603 0,311

9,000

3,405 -0,695 f0 ,738 0,811

1,946 -0,054 f0 ,615 0,081

2,433 $0,433 f0 ,615 0,70# - 1,216 +O,216 f0 ,465 0,46!

9,000

4,714 +01714 fo ,563 1,261

1,286 -0,714 fo ,471 1,511

1,714 -0,288 &0,471 0960'

- 1,286 +O,286 f0 ,356 0,80:

9,000

2,127 +(),la7 fO , l91 0,681

0 , 8 7 3 - ) ) )) ))

3,000 -


1912 ))

D

n ))

1)

B

))

))

n ))

TABLE 3. Segregation in Fs of AaHbCc-Fs-individuals (purple).

238 12 239 29 244 26 260 17 267 12 275 4 276 11 293 7 296 28 297 28 446 9

1 1 2 1 9

Total .................. Ratio .................. Found .................. Standard error, M, D/M, ..._.............

1913 224 )) 227 >) 231 )) 232 )) 253 P 255 )) 258 )) 260 )) 265 )) 266 )) 269 >) 270 D 274 n 281 )) 282

286 )) 290 1) 291 >) 302 9) 303 R 305 n 313 N 314

Deviation, D .........

Hercdlta. I.

Violet

183 27 27,428

f 1,530

0,280

39 10 36 9

22 29 19 12 11 27 32 18 28 10 7

32 12 30 30 22 18 22 19

f 0,429

4

- 2

2 3 2 4

18 3 2,698

-

- 0,902

f 0,655 0,461

6 4 7

3 2 2 1 2 4 6 2 3 1

3 1 4 4

-

-

6 8

6 5 7 2 2 6 6 6

-

9 5 7 7

17

l4 7 I 113 16 16,836

+ 0,836 f 1,341

0,608

26 12 24 7

12 21 11 7

15 20 22 21 18 5 8

23 5

20 1G

54 9 8,084

- 0,906 f 1,077

0,841

11 3

12 4 5 6 2 8

12 9

13 9 4 9 3

14 -1

10 5 5 2 6 4

2 2 3 4

Rose I

7 7

14 17

5 1

13 6

3 - 2 3

16 5 1

59 9 8,843

,& 1,077

- 0,157

0,146 I

12 5

15 4 5

10 8 4 8 3

10 4 7

4 9 5 6 9

11 9 (I

15

1

Total

8

41 63 40 41 29 18 22 21 70 55 27

427 64 64

94 34 94 24 47 68 42 32 46 63 83 54 60 31 22 81 27 70 6-1 47 38 53 59

G

82 HANS TEDlN

6 1 6 8 3 2 3 7 4 6 4 7 1 5 2

Year

23 I 61 18 54 7 21

21 62 6 33

11 55 16 38

1 1 2 1 3 1 4 I I I

1913

))

n ))

))

n ))

317 318 321 323 324 329 336

2 218

9 8,784

- 0,136

f 0,568

0,423

59 Total 1 9 1 2 ............ )) 1913 ............

4 1 5 35 93 445 1592 3 16 64 3,739 17,889 64 + 0,739 + 1,889 -

f 0,339 f 0,695 - 2,180 2,718 -

18 113 427

Suin total ............ Ratio .................. Found .......... .-. .... Deviation, D ......... Standard error, & DIM, ..................

18

26 10 32 11 16

633 27

i

25,447 - 1,553

f 0,792

1,961

816 27 25,864

- 1,136

f 0,703

1,616

7 2 4 7 4 4 8

203 9 8,161

- 0,839

f 0,658

1,504

257 9 8,147

- 0,853

f 0,495

1,723

218 I 93

Hose

445 1 1592

Total

277 9 8,782

- 0,218

f 0,496

0,440

111 3 1 "E 3,619 17,688 + 0,519 + 1,688

f 0,301 f 0,617

1,724 2,736

~~~~

I

1912 269 4 1 - 1 1 1 )) 1 270 10 4 1 - n 271 7 1 1 3 )) 280 19 8 3 5 )) 287 14 3 5 3

290 28 9 11 1 )) 453 14 2 8 1 ))

2019 64 64 -

- - - - - - -

TABLE 4. Segregation in Fa of AA13bCc-FI-individuals (purple).

)) I 454 Total .................. Ratio Found Deviation, 1) ......... Standard error, M,

..................

..................

DIM, ..................

- ' 5 4 1 1 1 1 - - 101 34 16

3 1 3,056 1,438 - + 0,059 - 0,673 + 0,056 + 0,438 -

& 0,m f 0,468 f 0,488 0,zso -

- 9 I 2; 9,079 2,027

0,133 1,224 0,120 1,610 -

8

6 15 12 35 25 49 25 11

1 78 16 16 -


D

))

>)

1913

D

D

11 8 2 1 - 37 242 3 6 2 - 21 261 14 4 5 3 284 11 7 5 1 - 24 297 40 3 11 6 - 60 308 13 7 13 4 - 37

?6 - 230 I i:

)) I 328 Total ... . .. ... ... ... ... Ratio ...... ...... ... ... Found .................. Deviation, 1) ......... Standard error, M, D/M, ..................

)) I 322 13 2 I 20 5 , - -

Total 1912 ............ I 101 27 I 34 I 16 I - 1 178

56 1 43 42 9 3 8,834 2,695 1,111

-- 0,160 - 0,405 + ::::: 1 - 0,111

f 0,493 f 0,888 f 0,388 f 0,211

0,337 1,044 1,183 0,465

)) 1913 ............ Sum total ..... . ... ... Ratio ...... ... ...... ... Found .................. Deviation, 1) .. . ... ... Standard error, M, D/M, ... ... ... ... ... .. .

- 259 - 16

16 -

- -

214 9 8,934

- 0,086

f 0,980

0,174

56 I 18 I - I 259

3,295 1,245 - 16

f 0,299 f 0,185 - 0,987 1,324 -

69 3 2,526

- 0,414 f 0,289

1,586

242 1 8 3 )) 14 4 D 26 5

1912 1 250 253

D 265 9 4 D 274 25 9 )) 277 16 5 D 295 16 2 )) 299 48 13 D 447 23 7

- 2 13 - 12 30 - 17 48 - 3 16

- - 5 39 - 3 24

- - 11 29 - 18 79 - 9 39

- - - -

-

- -

Total ......_........... 185 52 - - 80 Ratio .................. 9 3 - 4 - Fount1 .,.. ..,............ 9,338. 2,625 I - I - 4,037 Deviation, 1) ......... -i- 0,388 - 0,375 - f 0,031 Standard error, M, f 0,440 j= 0,351 - 1 1 f 0,389 D/M, .................. 0,758 0,008 - - 0,095

317 16 16 - - -

84 HANS TEDIN

1912 n R

Year

1

249 266 457

Fieid- number

2

- 1 ; 5

17 9 6

- -

226 233 238 268 255 277 278 279 283 288 300 301 328 331 332

25 - 1 ; - 7 18 18 -

rota1 .................. Ratio .................. Found .................. Deviation, D ......... Standard error, M, D/M, .................. rota1 1912 ............

)) 1913 ............ Sum total ............ Ratio .................. Found .................. Deviation, D ......... Standard error, M, DIM, ..................

- - 10 1 l l - 7 1 -

Purple

3

4 I ::

27 23 45 8 11 49 29 36 21 13 17 9

13 10 1'5

350 9 8,531

--

- 0,463

f 0,310 1,494

185 350 535

9 8,798

- 0,101 f 0,254

0,796

Violet

4

7 3 9

17 1

18 12 15 6 5 2 5 4 4 8

116 3 2,829

- 0,171 f 0,244

0,701

52 116 168

3

- 0,238

f 0,200 1,190

2,762

Rose Light purple White

7

19 8 9

22 5

34 14 21 11 10 3 8

10 7 9

190 4 4,634 + 0,634

f 0,271 2,339

80 190 270

4 4,440

f 0,440

f. 0,222 1,982

Total

53 34 63 71 17

101 55 72 38 28 22 22 27 21 32

656 18 16 - - -

317 656 973

16 16 - - -

TABLE 6. Segregation in FJ of AaBBCc-Fr-individuals (purplej.

8

TllE INHERITANCE OF FLOWER COLOUR IN PISUM 85

4,414

+ 0,414

f 0,743

0,557

9 14 16 3 2

15 7

11 10 14

1 1 2

FQund .................. Deviation, D ......... Standard error, M, D/M, ..................

16 - - -

26 49 47 22 14 50 30 37 24 56

225 228 229 236 237 267 296 304 306 31 1 319

109 4 4,663

f 0,WS f 0,358

1,852

21 109 133

4 4,616

$- 0,616

f 0,3as 1,907

Total .................. Ratio .................. Found .................. Deviation, 1) ......... Standard error, M, D/M, .................. Total 1912 ............

)) 1913 ............ Sum total ............ Ratio .................. Found .................. Deviation, D ......... Standard error, M, D/M, ..................

374 16 16 - - -

87 374 461 16 16 - - -

Purple

D

u ))

3

289 292 316

9,011 + 0,011

f 0,861

0,013

14 29 23 10 11 27 17 22 13 29 10

205 9 8,770

- 0,230

f 0,410

0,561

49 205 254

9 8,816

- 0,184

f 01370 0 , w

Violet RoBe

5

2,675

- 0,425

f 0,670

0,634

3 6 8 I) 1 8 6 4 1

13 1 60 3 2,567

f 0,323

1,34 1

- 0,453

14 60 74 3

-

2,568

f 0,291

1,485

- 0,432

Light purple

6

White I Total

TAISLE 7. Segregation in Fs of AAIJb(:(:-F1-individiials (purple).

1 1 2

86 HANS TEDIN

1912 1)

n n B

))

243 247 255 279 294 450

Ratio .................. Found ...... .......... Deviation, D ......... Standard error, M, D/M, .................

3 3,150 + 0,130

f 0,120

1,083

1 0,870

- 0,130 f 0,120

1,083

4 4 -

TABLE 8. Segregation in Fs of AABBCc-FAndividuals (purple).

3 5 8

13 11 36 10 23 9

102 3 3,114

+ 0,114

0,766

f 0,111

35 15 46 25 42 16 49 20 32 43

1 5 4 7 9 3

29 1 0,886

f 0,161

20 3 9 8

21 3

16 5

10 10

- 0,114

0,766

14 16 40 17 32 12

131 4 4 - - -

55 18 55 33 63 19 65 25 42 53

1913 n n n ))

n B n n n

247 251 271 272 293 327 335 346 364 382

105 1 0.981

- 0,019

OJ26

f 0,081

29 105

428 4 4 - - -

131 428

rota1 .................. Ratio ...... !. .......... Found .................. Deviation, D ......... Standard error, M, D/M, .................. rotal 1912 ............

n 1913 ............ Sum total ............

323 3

+ 0,019 3,019

f 0,084

0,226

102 323 425 134 559

THE INHERITANCE O F F L O W E R COLOUR IN PISUM 87

1912 Y

))

Y

Y

251 264 451 455 456

19 7

29 23 5 5

11 13

60 37 74 69 38 17 34 75

135 1

+ 0,142

1,142

f 0,080

1,115

21

473 4 4 - - -

87

Total

8 5 1 6 1 7

- - - -

- O,C41 - f. 0,013 -

0,582 1 - 1 -

Ratio .................. Found .................. Deviation, D ......... Standard error, M, DIM, ............ :. ....

3 3,041

f 0,041 f 0,073

0,582

TABLE 9. Segregation in Fs of AaBBCC-Fn-individuals (purple),

7 1 8

2 1 :; f l 9

8

11 I 38 21 I 87

3

13 13 7 6

27 66 3 3,035 + 0,055

f 0,186

O,l8i3

41 30 45 46 33 12 23 62 46

1 0.965

- 0,035

f. 0,188

0,188

241 244 252 280 294 298 307 309 310

rota1 .................. Ratio .................. Found .................. Deviation, D ......... 3tandard error, M, D/M, .................. rota1 1912 ............

n 1913 ............

338 3 2,858

f 0,080

66 338

- 0,142

1,115

sum total ............ Ratio ..................

404 3 156 1 I 4

88 HANS TEDIN

.................. Found Deviation, D ......... Standard error, M, D/M, ..................

1,114 4 2,881 - - - - 0,114 - - - + 0,114 -

- f 0,073 - f 0,073 - - 1,662 I - - I - 1,562 I -

TABLE 10. Segregation in F2 of AabbCc-Frlindividuals (violet).

I 2

240 252 %7 261 262 263 2tx2 286 288 291 292 2D8 301

rota1 .................. Ratio .................. Found .................. Deviation, D ......... itandartf error, M, DjM, ..................

344 348 352 354 358 385 369 373 374 376 377 378

3 =

4

18 38 14 42 14 3

?5 26 15 11 15 29 31

281 9 9,669

+ 0,680

f 0,368

1,818

23 31 11 29 10 34 14 16 19 16 36 12

6

5 8 8

14 4

11 6 5 1 6 5 5 8

89 3 3,062 0,062

f 0,290

0,214

9 11 8

11 5 8

13 6 5 7 6 7

7

10 18 3 9

10 1 4 5 5 9 4 9 8

95 4 3,268

f 0,321

2,277

- 0,731

8 8 5

10 3

19 9 8

27 6

21 6

8

33 64 25 65 28 18 35 36 21 26 24 43 47

465 16 16 - - -

40 50 24 50 18 61 36 28 51 29 63 25

THE INHERITANCE OF FI.OWER COLOUR IN PISUM 89

- - - - - -

- -

323 9 8,571

- 0,429 f 6,323

1 , a n

281 323

- - -

604 9 9,049

- - - - - -

- - -

1 69 3 3,059 + 0,059

f 0,117 0,504

3 6 36 19

Light purple

G

White

7

Total

8

Purple Violet

3 1 4

Rose

5 -

1 1 2

1913 ’ 380 )) 1 385

Total .................. Ratio .................. Found .................. Deviation, D ......... Standard error, M, D/M,, .................. Total 1912 ............

)) 1913 ............

96 32

12 5

28 11

113 3 2,098

- 0,002 f 0,254

0,008

89 113

167 4 4,431 + 0,431

+ 0,282

95 167

1,528

603 16 16 - - -

465 603

Sun] total ............ Ratio .................. Found .................. Deviation, D ........ Standard error, M, D/Mi ..................

202 3 3,oaa + 0,026 0,138

f 0,191

262 4 3,925

- 0,075 f 0,212

0,354

1068 16 16 - - -

TABLE 11. Segregation in F8 of AAbbCc-Fr-individuals (violet).

5 8 7 1 2

241 254 283 284 285 289

5 7 14 10 12 4

22 16 63 53 49 18

Total .................. Ratio .................. Found .................. Deviation; D ......... Standard error, M, D/M, ..................

52 1 0.941

f 0,117 0,504

- 0,050

10 22 9 2 4

11

221 4 4 - - -

46 58 28 12 10 41

340 341 349 360 362 366

1913 b

1)

))

1)

1)

90 HANS TEDlN

))

N N

Y

n I)

N

D

1)

_- Purple

3

339 343 347 353 355 37 1 372 389 390

Light purple

ti --

Violet

4

Rose

5

White

I

Total

8 1 1 2

1913 1 370 )) 375

Total .................. Ratio .................. Found .................. Deviation, D ......... Standard error, M, D/M, .................. Totai 1912 ............

N 1913 ............ Sum total ............ Ratio .................. Found .................. Deviation, D ......... Standard error, M, DIM, ..................

28 14

10 6

38 20

1 79 3 2,8SO

- 0,110

f 0.10s

1,560

1 69 179

74 1 1,110

f 0,109

52 74

f 0,170

1,560

253 4 4 - - -

22 1 253 474

4 ? - - -

348 3 2,957

- 0,oea f 0,080

0,788

126 1 1,003

f 0,080

0,188

f 0,063

TABLE 12. Segregation in Fa of AabbCC-Fg-individuals (violet).

1 1-7 7

9 4 6 8

28 11 12 14 12 22

126 1 1,400

f 0,400

f 0,091

4,396

18 20 16 19 68 17 47 36 48 71

360 4 4 - - -

TI-IE I N H E R I T A N C E O F F L O W E R COLOUR I N PISUM 91

14 225

TABLE 13. Segregation in FB of AaUbcc-F%-individuals (rose).

50 1 772

--

Year

1

- Field-

number

2

302 304 305 306 310 314 315 31 7 319 320 32 1 325 326 327 328 333 338 339 340 342 345 346 348 349 359 362 365 368 370 37 1 372 373 375

rota1 .................. Ratio .................. Found .................. Deviation, D ......... Standard error, M, D/M, ..................

-__

Purple

3

Violet

4

Hose

s

5 12 26 2 7 24 10 3

17 14 13 14 6

19 10 14 10 4

10 15 17 16 5 3 4

12 14 10 8 7

38 10 25

404 9 8,m

f 0,986

- 0,627

2,192

Light purple

G

2 4 6 1 2 4 6 2 3 5 6

10 1 3 7 4 2 2 2 4 4 1 1 9 4 8 4 2 1 1

12 9

11 143

3 2,964

- 0,036

f 0,226

0,lSCr

White

7

2 4 2 1 3

12 7 1

11 4

21 7 2

10 6 8 8 2 4 5 3 8 3 9 4

12 15 3 2 1

19 12

Total

8

9 20 34

4 12 40 23 6

31 23 40 31 9

32 23 26 20 8

16 24 24 25 9

21 12 32 33 15 11 9

69 31

4 4,663 + 0,663

f 0,249

2,663

16 16

92 HANS TEDlN

TABLE 11. Segregation in FI of AABbcc-FAndividunls (rose).

Year Light purple White I Total Field-

number

2

Hose

5 6 1

39 6

10 4

10 10 9 7

12

14 4 2 8 5 5 2

11 4

53 10 12 12 15 15 11 18 16

303 318 322 323 330 343 361 364 366

rota1 .................. Ratio ................?. Found .................. Deviation, D ......,... Standard error, M, DIM, ..................

107 3 2,sra

- 0,358

f 0,136

2,832

55 1

f 0,958

f 0,138 2,832

1,368

TABLE 15, Segregation in FS of AaUBcc-Fn-individuals (rose). =

8 2 3 4 5 6 7 1

308 324 329 331 332 335 336 347 351 353 360 374

3 4 8 1 6 9 6 7 2 5 5 7 63 1 1,086 + 0,086

f 0,114 0,754

11 26 30 12 17 19 15 21 16 12 12 41

232 4 4 - - -

8 22 22 11 11 10 9

14 14 7 7

34 169

3 2,914

- 0,086

f: 0,114 O,?M

rota1 .................. Ratio .................. Found .................. Deviation,, D ......... Standard error, M, D/M, ..................

T H E INHERITANCE O F FLOWER COLOUR IN PISUM 93

TABLE 16: Segregation in Fs of Aabbcc-Fa-individuals (light purple).

rota1 .. . . . . . . . . . . . . . . . .

376 379 381 382 384 385 386 387 388 389 390 391 395 397 398 400 401 402 403 405 407 408 409 410 414 416 417 419 420 421 422 43 1 432 433 435 437 438 439 440

Purple

3 4 1 5

I I

- - - I -

I I - -

- I - - I -

Light purple

6

38 16 8

16 13 29 22 50 17 10 5

18 + 4 34 8

34 17 26 28 20 14 17 17 22 11 4

13 2

18 34 28 37 7

22 36 13 28 43 10

789

White

16 14 7 10

2 8 2

16 13 12 4 6 5 9 7

28 4

11 10 12 10 9 -4 7 2 4 4 4 4

14 7

10 5 7 4 3

13 22 6

335

Total

8

54 30 15 26 15 37 24 66 30 22 9

24 9

43 15 62 21 37 38 32 24 26 21 29 13 8

17 6

22 48 35 47 12 29 40 16 41 65 16

1124

94 HANS TEDlN

.................. Ratio -

Deviation, 1) ......... - Fount1 .................. - 1 - -

- Standard error, M, - - D/M, ................. I - -

Year

3 1 4 - 2,808 1,ioa 4 - - OJoa + 0,192 -

-

- & 0,062 f 0,062 - - 3,602 3,691 -

Hose

Year S e g r e g a t i o n i n Fa Field-

Total n u m l w Purple 1 Violet I .nose I I White

TABLE 17. 01001, AAbbcc, light purple 9 X aaBBcc, white d (The white from FS of a cross quite like cross I table 1).

FI AaBbcc, rose.

11

1911 336 338

)) 339 )) 340 )) 344 )) 315 h 346 )) 347 )) 348

350

)) 353 Total .................. Ratio ..................

1 351

Found ..................

Standaril error, M, Deviation, D .........

D/M, ..................

14 i 44 8 1 12

1 1 52 - 32 ' 9 - - 14 6 7 27

- 35 5 14 54 - - 24 3 11 38 - - 25 7 12 44 - - 35 8 13 56 - - 20 4 6 30 - - 13 2 1 16

13 12 51 - - I ;: 9 9 32

- 268 77 111 456

= I : - -

- : I -

- -

- : : I 9 3 4 16 - - 9,404 2,702 3,804 16 + 0,401 - 0,208 - 0,106 -

j= 0,372 & 0,202 f. 0,824 - - 1,086 1.021 0,327 -

TABLE 18. 01001, AAbbcc, light purple 9 X aabbcc, white (from R of cross I.table 1) d.

F1 Aabbcc, light purple. _ _ _ _ _ 1 1 2 1 3 1 4 1 5 1 6 1 7 1 8

104 74

I 1913 )) 1 ' 391 392 ' I - - I - - 1 - 1 4 0 ' -

T H E INHERITANCE OF FLOWER COI.OUH IN PISUM 95

42 10

2 6 / - 1 8 8

43 1 1 1 - I 19 8

Year

12 12 17

- - - -

1

119

8,774 9

- 0,226

Field- number

2

- - 53 217 i 4 : I - l 4 1 1 6 - 3,318 - 1 3,908 I 16 - -/- 0,318 1 - - 0,082 - - 1

393 394 395 396 397 398 399 400 401' 402 40'1 404

!%: 407 408 409

I'otal .................. Ratio .................. Found .................. Deviation, 1) ....... :. Standard error, M, DIMk .................

Purple

I S e g r e g a t i o n i n F2

Violet

7 25 16

85 22 53 23 62 13 47 17 63 18 31 11 15 5 38 16 45 26 36 15 51 25 34 13 40 20 87 23

9J5 330 3 1

71 I ;;

2,957 1,013

- 0,043 + 0,043 f 0,048 f 0,049

0,878 0,878

23 37 93

107 76 75 64 81 42 20 54 71 51 76 47 60

110 12135

4 4 - - -

TABLE 19. Cross I. White, aaBBcc 9 X 0401, AABBCC, purplc 6. (The white froiii Fz of cross I table 1).

)) 11. The same whitc Q XO351, AABBCC, purple 6. FI. In both cases AaBBCc, purple.

' 1 1 2 1 3 1 4 l 5 1 6 1 7 1 8 1

1913 I I 461 )) 462 )) I 463 )) 464

Total .................. Ratio ... .:. ............ Found ................. Deviation, D ..........

96 HANS TEDIN

f 0,589 O,4lB

46 26 50 21 41

184 9 9,058 + 0,058

f 0,440 0,132

S e g r e g a t i o n i n Fa I

- - - - - - - - - - - - -

Field- number

Year

1913 I1

))

))

1)

))

465 466 467 468 469

Rose

3 1 4 5 6 1 7 1 8

- f 0,470 - - 1 0,198 1 -

f 0,424 0,750

26 16 9

11 10 72 3 3,545

f 0,545 f 0,346

1,575

14 8

18 14

86

77 46

50

15 I 66 69 I 325 4 3,397

- 0,603

f 0,384 1,570

16 16 -

TABLE 20. 0142, AAItBcc, rose Q X aaRBCC, table 1). d.

FI. AaBBCc, purple.

white (from F1 of cross I

1 2 3 7 8 5

445 446 447 448 449 450 452 451 455 457 458 459 460

............

78 42 52 63 43 32 25 35 48 32 29 56 63

124 71 91

115 91 55 52 71 87 72 62 91 98

1080 16 16 - - -

15 13 14 20 21 7 9

17 1% 13 10 18 14

189 3 2,800 0,200

f 0,100

1,053

31 16 25 32 27 16 18 19 21 27 23 17 21

293 4 4,341

0,SU

1,616

f 0,211

598 9 8,859 0,141

f 0.242 0,583

Ratio .................. Poiind .................. Deviation, D ......... jtandard error, M, D/M, .............. ;...

T H E INHERITANCE OF F L O W E R COLOUR I N PISUM 97

LITERATURE CITED.

1; TSCHERMAK, E. v. 1902. Ueber die gesetzmlissige Gestaltungsweise der Misch-

2. -

3. -

4. -

linge. 1904 a. Die Theorie der Kryptomerie und des Kryptohybidismus. Beih. z. Bot. Centralblatt. Bd. XVI, p. 11-35. 1904 b. Weitere Kreuzungsstud. an Erbsen, Levkojen u. Bohnen. Zeitschr. f. d. landw. Versuchsw. in Oest. p. 533-638 1912. Bastardierungsversuche an Levkojen, Erbsen u. Bohnen mit Riicksicht auf die Faktorenlehre. Bd. VII,

Zeitschr. f. d. landw. Versuchsw. in Osterreich p. 781-861.

Zeitschr. f. ind. Abst. u. Vererbungslehre. p. 81-234.

5. WHITE, 0. 1917 a. Inheritance studies in Pisum IV. Interrelation of the genetic factors of Pisum. Journal of agricultural research v. XI, p. 167-190.

6. - 1917 b. Studies of inheiitance in Pisum 11. The present state of knowledge of heredity and variation in Peas. Proc. Amer. philos. SOC. v. LVI, p. 487- 588. Unfortunately, I have not been able to consider this paper as only part

7 LOCK, R. H. 1908. The present State of knowledge of heredity in Pisum. Ann.

8. KAJANUS, B. u. BERG. S. 0. 1919. Pisum-Kreuzungen. Arkiv fiir Botanik. Bd 15.

1 and 2 of volume 56 have come to Sweden.

of bhe Royal Bot. Gard. Paradeniya. v. IV.

N:o 19, 18 p.

Pt. 111, p. 93-111.

Hereditus I . 7

Documents

THE INHERITANCE OF FLOWER COLOUR IN PISUM