Historic, archived document

Do not assume content reflects current
scientific knowledge, policies, or practices.

Issued January 13, 1912.

U. S. DEPARTMENT OF AGRICULTURE,

FOREST SERVICE—Circular 196.

' HENRY S. GRAVES, Forester.

THE INFLUENCE OF AGE AND CONDITION OF
THE TREE UPON SEED PRODUCTION
IN WESTERN YELLOW PINE. |

BY

G. A. PEARSON,
In Charge, Fort Valley Experiment Station.

Tests on western yellow pine seed

Germination 2» =.5 2.22: 5.

MAPICS ON eto eee

Application of conclusions
[Oir. 196}

CON TEN TS.

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THE INFLUENCE OF AGE AND CONDITION OF THE TREE UPON SEED
PRODUCTION IN WESTERN YELLOW PINE.

TESTS ON WESTERN YELLOW PINE SEED.

The influence of the character of the mother plant upon the quality
and quantity of seed produced and upon the character of the offspring
is a factor which is well recognized in plant breeding. In forestry
this factor should be considered not only in collecting seed for arti-
ficial propagation, but also in the selection of trees to provide for the
natural regeneration of cut-over areas. Im selecting seed trees the
forester must ask himself: ‘“‘ Which trees will produce the best quality
and the greatest quantity of seed?” and finally, ‘‘ Which will produce
the most desirable seedlings?’ Secondary, though very important,
considerations which should govern the forester in his selection of
trees to provide for the future stand after cutting are wind firmness
and merchantable deterioration or appreciation.

This circular deals primarily with the conditions affecting the
germinative quality of the seed. The influence upon the quantity
of seed, though of great importance, will be treated only as a second-
ary topic, because the data on this phase of the subject are incomplete.

The species studied was western yellow pine (Pinus ponderosa).
The locality was near Maine, Ariz., on the Tusayan National Forest,
elevation about 7,200 feet. In November, 1909, 100 samples, each
consisting of from about one-tenth to 1 bushel of cones, according to
the amount available, were collected from 100 trees representing
different ages, sizes, and conditions of soundness and general health.
The cones were collected in connection with a logging operation, which
made it possible to determine accurately the age and soundness of
the tree. In the case of young trees which were not felled the age and
soundness had to be estimated;.but since the number of such trees
was small and nearly all of them were thrifty, the errors in these esti-
mates will not be great enough materially to affect the results. In
each case the sample was made up of cones from all parts of the
crown. Careful data were taken on each sample, according to the
outline on page 4.!

1 The seed samples and accompanying data were gathered by Forest Assistant Harold H. Greenamyre.
16333°—[Cir. 196]}— 12 (3)

gies es ee ae paper ecegiyge tie Sy ey ARC Yield of cones (bushels)..002 055. Ae oso
Heieiihi(feety oc aoe eke ce eee Gontlition of canes.) 002 Soo ee ae
Diameter breasthigh (inches)...........- | Number of cones in sample.............
Ace (years) occ: 2 eee ane aca Dry weight sample (pounds)..........
Clear Tenethe Ws ae Rin ee ie eee Weight seed (ounces)..............--2.--
Condition of boley 7s. Sy eeuhe nee eee Site:

Condition eh Craw. os) oe ae eee Exposure and slope... .-..8 22.2022
Fire gear? - sz acehe epee: eine. cee: eee Poth. cpa's oy eres: Saree
Spikerbawe [AY bk RRO RARE oa. oe RYOIChEN REV Le eos ee ee oem ee
Other peculiarities: 47485) dbus en RESIS BD ERE E SS ee Se

TABLE 1.—Comparative germination of seed of western yellow pine—Coconino Experi-

ment Station.

I |
uration of te ys). _
Durat test (days) howe Aver
| i || age asi
| 2 > | rield amount
Tree class. eye Oe Ua hae fe Basis.|| of |Basis | of seed
per
ee pound
Average germination | Hae of dry
(per cent). cones
| | | 1)
} | Trees. Bush. |Trees.|| Ounces.
Yellow pine............ | 42 | 58 | 61/64/67) 68) 60]) 1.8] 55] 13
Baek ack sauce 5S | 69 | 72 |,74 75 | 76 38 | 1.0 8 7
Healthy yellow pine...) 41} 55 | 57 | 61| 63 | 64; 28 || 18] 27 1.2
Healthy blackjack ... ..| | 53.| 71 | 74] 76 | 77) 78). 34]) LO St A Aeb
Unhealthy yellow pine.| 43 | 58 | 65/69/70) 71) 34|| 17! 30 14
Unhealthy blackjack ..| 44 | 55 | 57 | 59 | 60 | 61 | eee [ae 19
Open'stand!;:1). 2 Y22. -| 46 | 62 | 65 | 68 | 70 | 70 TA Nak ed 51 1.4
Medium stand!........ | 41.157] 61|65|68]/69| i8|| 241] 13 | 1.5
Dense stand!.......... | 56| 77 | 78 | 79 | 791 79 9 || My 4] 18
By decades:1 | |
Were artes 63 | 78 | 80 | 82 | 83 | 84 ep t ace 3] #15
To eee 68 | 84 | 86 | 87 | 87 | 87 dhe a ldeilced 2 1.0
BED we tL ve ee Poe al 55 | 68 | 70 | 71 | 73 | 73 7 8 6 1.5
A ecto = ed ech 53 | 68 | 70 | 72 | 74 | 74 a oes Be 6 1.3
Be 15 GB, oe 36 50 | 54) 58) 6L/ 62} 11 |) 17) 10 1.4
°C pie Soll RS ie 48 63 | 69 | 75 | 77 | 77 4 1.7 4 1.2
meta ye eel | 85. | 74 81 | 82 | 82 | 82 1}} 40 Ly ae!
BiGh poate’. els 133.155|58| 61] 64\66| sil 21] sil 3
Sp AS # ee | 99 | 42 | 45 | 47] 49 | 50 BW iho 5 119
7 RR So SRG SET 56 |. 77 | 78 | 82 | 83 | 84 2 1.5 1 1.0
MNO Oe 48 | 56 | 61|66|72|74| 2 CS BRS ae
2 Sa ae 54 | 86 | 86 | 88 | 88 | 88 1 1.5 1 | 2.0
72, TO MAR NS 3 ae A a | a PE 2 4 Sle PT ce en a ee eS Ee FAS. Aleeacens e
Otte. fe et a Se ae a a Hee | Pee eeaes | ee | WLI IE
Pye he CO Re 20 | 24 | 26 | 33 | 38 | 39 2 Rie PAS
SOs ek. gi | 48. Soh en epitaes | 61 2 | 6 2 1.5
pms les nee ONT ee Pe aes eae eae de: Re: OO
Tie FG ere a 36 | 53 | 57 | 62 | 63 | 63 1 | 8 A BSR
SATEEN. Be | 3) 7] 8} 14] 22 | 36 1 2.0 1 1.3
2 REN 38 | 79 | 80 | 81 | 83 | 83 1 2.0 1 | .9
SAU MERE! 286 | 26 | 50 | 55 | 62 | 70 | 71 3 ie 2 1.4
2A cee SE | 70 | 80 | 82 | 83 | 83 | 83 2 3.3 2 2.0
aaa ELT FOLLAL SORT EOE Ey eee | AES DY Hg RR
2 MERI OAT oS (ET EE ENS Raabe vege) Rae Fie Ta ec tome eo
7 (ROD Bokeh ta} 12! 15 | 16 | 18 | 24) 27 | 1 2.5 1 1.2
5 ERG SER | 61 | 70 | 72 | 74 | 75 | 75 2 3.0 Oe | ieee. be
ASTER: Ph 43 | 76 | 77] 78 | 78 | 78 1 2.0 1 9
Below 140 (diameter
breast-high 14 inches
and below) ?........- 52 | 71 1°73 | 77.1) 784 81 “2,11 Pan ee se 1 RS Speer
PaN-PS01 Pics re he Ue | 45 | 61 | 65 | 69| 70 | 71} 36 1.6| 49 183
EP AM aces a | 37 | 52 | 54 | 58| 62|63| 16 | 20) 15 || 13

1 All trees.
2 Blackjack, not cut. Age estimated.

Aver-
age
num-
ber of | Basis.
|: seed
per
pound
Trees.
12, 065 62
12, 624 38
11,362 28
12,305 34
12,644 34
15,352 4
12, 223 73
| 12,498 18
| 12,294 9
Ratan | aoe
|
\ |
|
| ae. wRE
|
Pes) 6) 2h}:
13, 256 AL
11, 652 63
13, 447 16

3 Includes 10 ‘‘ blackjack” 15-22 inches diameter breast-high and 1 ‘‘yellow pine” 17 inchcs diameter

breast-high whose exact age was not determined.
[Cir. 196]

Oo

Table 1 gives a comparison of the different classes of trees with
reference to germination and yield.

The cones were air dried. Each sample was placed in a 2-bushel
sack immediately after collecting and kept there through the entire
process of drying. The amount placed in each sack was in no case
more than 1 bushel, though by the time the cones were fully opened
the volume had about doubled. The drying process, which involved
daily turning of the sacks, required about six weeks. No rain or snow
fell in the meantime. The seeds were extracted by first shaking the
cones violently in an iron tub and then striking each cone separately
against the side of the tub. They were cleaned by rubbing between
the hands, and later allowing the wings to blow out by pourmg the
mass slowly from one vessel to another in a light breeze, care being
exercised to prevent the light, hollow seed shells from blowing out,
because obviously these should be considered in the germination tests.

The germination tests 1 were made at Washington by planting 200
seeds from each sample in a sand flat and subjecting them to a hot-
house temperature of 60° F. The duration of the test was 52 days,
during which period counts were made every two or three days.

The terms “‘blackjack” and ‘‘yellow pine” will be used in this
circular to distinguish two forms of the species Pinus ponderosa.
“Blackjack” is the local name applied to young, vigorous trees
characterized by a dark, almost black bark. ‘‘Yellow pine”’ is the
local name applied to the older trees characterized by a yellowish or
reddish brown bark. Lumbermen generally regard the two forms
as distinct species, but the only difference is that due to age.

From Table 1 the following deductions may be made:

GERMINATION.

Seed from young trees shows a higher germination than seed from old,
mature, or overmature trees. The 38 blackjacks used in this test
give a germination of 76 per cent, against 68 per cent for the 60 yellow
pines. A comparison of 20 trees near Cliffs on this Forest in 1908
gave a germination of 68 per cent for yellow pine, against 80 per cent
for blackjack. Classifying both blackjack and yellow pine accord-
ing to age, by decades, the germination per cent is found to fall
with the increase in years. This decline is, however, by no means
regular. For instance, 5 trees 220 years old show an average germi-
nation of only 50 per cent, while 2 trees 390 years old show an aver-
age of 75 percent. Dividing all trees, both yellow pine and blackjack,
into three age classes, namely, below 140 years (8 to 14 inches diam-
eter breast-high), 140 to 250 years, and 260 to 400 years, the average
germination per cents are found to be 81, 71, and 63, respectively.

: 1 Conducted by Mr. N. H. Grubb, Office of Silvies.
(Cir. 196}

6

Certain injuries decrease while others increase seed quality. A com-
parison of healthy and unhealthy trees shows the following germina-
tion per cents: Healthy yellow pine, 64; unhealthy yellow pine, 71;
healthy blackjack, 78; unhealthy blackjack, 61. In classifying the
trees with respect to health, those affected by heart rot, insects,
mistletoe, dying top, fire scars, and suppression are classed as ‘‘un-
healthy,’ while those showing no indications of disease or injury are
classed as ‘‘healthy.”” These terms mean but httle, however, unless
we inquire into the actual condition of the tree. Grouping the 34
trees classed as unhealthy according to the nature of the injury, the
following interesting comparison is derived: :

TABLE 2.—Comparative quality of seed from healthy and unhealthy trees.

| Healthy. | Unhealthy.

| |

Heart Spike | Fire sa, a Saft
¥ ote fae rot, yel- | top, yel- | yello beste, It efblacke
Pp | Je low pine. | low pine. ne cA
ERB ie ee) |
Germination......... per cent. .| 64 78 67 79 79 37 51
BSIGP Se Rene Ae 2 Ae trees. | 28 34 10 | 7 15 3 3

Yellow pine affected by heart rot shows a germination 3 per cent
higher than that of sound, healthy yellow pine. This difference, based
upon only 10 trees, is hardly sufficient to establish the conclusion
that heart rot increases germinative quality, but at the same time
tends to show that at least the seed of such trees is not inferior to
that of sound trees. The smallness of the effect is probably due to
the fact that the rot works upon the heartwood exclusively and
does not directly attack the vegetative life of the tree. If the rot
should become sufficiently advanced to reduce materially the vitality
of the tree, it is probable that the quality of the seed would be im-
paired and the quantity diminished. On the other hand, the fungus
in its earlier stages may act as a stimulus, as in the case of ‘‘spike
top”’ and fire scars, considered in the paragraphs following.

Spike tops show a germination 15 per cent higher than that of the
healthy yellow pine. ‘‘Spike top” is the common term applied to trees
with dead leaders, generally caused by some mechanical injury, most
frequently lightning. The wound is usually followed by a fungous
attack. The leader dies back from year to year, and as the upper
branches drop away a straight shaft is left projecting above the
living portion of the crown. The injuries causing spike top evi-
dently act as a stimulus upon seed production. This peculiarity—
apparently nature’s provision for the perpetuation of the species—
although not fully understood, is a matter of common observation.

[Cir. 196]

7

It is in accordance with the same natural law which governs sprout
growth after a basal burn, and the production of adventitious shoots
after an injury on the stems or branches of certain broadleaf trees.

Yellow pine marked by fire scars shows a germination 15 per cent
above that of healthy yellow pine. Fire scars are confined almost
entirely to the base of the tree. They begin as a small burn in the
bark and cambium, but increase in size and depth with recurring
fires, until finally they sometimes extend 5 feet or more up, and one-
half or two-thirds through the stem. Trees thus affected generally
bear no marked signs of reduced vitality except in advanced stages,
although possibly measurements might show a diminution in growth.
The direct effects of the burn are a reduction of the water supply of
the crown of the tree because of the destruction of more or less of
the water-transporting elements, and at the same time a reduction
of the food supply of the roots through the destruction of some of
the elements bringing elaborated food material from the crown to
the roots. It is evident that the food supply of the roots will suffer
more than the water supply of the crown, because the elements
bringing down the food, being nearer the surface, are more readily
destroyed by the fire. The eventual result will be a reduction of the
food supply of the entire tree, due to a reduction of the amount of
water: which the tree is able to absorb. In spite of this fact, burns
appear to stimulate seed production, as does the injury causing
spike top.

Mistletoe-infected blackjack shows a germination 17 per cent below
that of unaffected blackjack. Although mistletoe stimulates growth
in the immediate region of the parasite, causing the abnormal devel-
opment of certain branches, the excess of carbohydrates produced
by the additional foliage of the mistletoe-stimulated branches is used
by the parasite itself, so that the final result is a drain upon the tree.
It seems highly improbable that mistletoe would ever act as a stimulus
upon seed production, because its action is insidious, slowly sapping
the vitality of the tree instead of causing injuries in the nature of
a shock, as is the case with burns or other mechanical injuries.

The effect of suppression 1s not clearly demonstrated. Of the two
yellow pine included in our data, one has a germination per cent of
92 and the other only 14.5, the average of the two being 53, or 11
per cent below that of normal yellow pme. The only blackjack
tested shows a germination of 90.5 per cent, or 12.5 above that of
the average healthy blackjack. On account of the great variation
between the individual trees cited above, it is impossible to predict
the result of averaging a large number of trees. One would naturally
expect to find an inferior quality of seed on a suppressed tree, but
this would vary with the degree and period of suppression. It will
also be counterbalanced to a certain extent by those factors which

[Cir. 196]

8

tend to better the quality of seed in dense stands, such as more
favorable conditions for pollination and the moderating effect of
close crown cover, described in the following paragraph:

The highest quality of seed 1s produced in dense stands. The germi-
nation for open, medium, and dense stands is 70 per cent, 69 per
cent, and 79 per cent, respectively. The difference between the open
and medium stands is too small to establish any conclusion one way
or the other; but the difference in favor of the dense stand is very
pronounced. At first thought one might expect to find the highest
quality of seed in open stands where light conditions are most favor-
able. There are, however, a number of factors which tend to explain
why the contrary is true. First, it is obvious that the denser stands
afford the best conditions for pollination; secondly, the moderating
effect of a close crown cover upon temperature and evaporation
would favor seed development, especially in the early stages. Many
young cones die during the winter and spring, probably as the result
of the unseasonable freezes and the drying winds which are very
prevalent. Flowers suffer in the same way. Instrumental observa-
tions at the Fort Valley station show that the minimum temperatures
under a fairly close canopy are commonly from 3° to 17° higher, and
the wind movement and evaporation from 50 to 100 per cent less
than in the open.

That the age and condition of the tree apparently have no mate-
rial effect upon the rate of germination is evinced by the fact that
the ratio between the germination per cents of different tree classes
remains practically constant throughout the period of the test.

YIELD.

No accurate data with regard to the absolute yield in seed are
available, but a rough measure of this yield is expressed by the
preduction of cones given in the preceding table. It should be
remembered that, in comparing yields with a view to determining
the influence of the character of the tree, it is necessary to consider,
in addition to the condition of soundness and general health, also the
crown development of each tree, a feature which is determined
largely by the density of the stand. Moreover, it is to be taken for
granted that, other things being equal, the yield will increase with
age until a certain age limit is reached. An exact study of the -
factors influencing both germination and yield is much more com- .
plicated than one in which germination alone is considered. Since
this study was undertaken with the primary object of determining
the conditions affecting germination, the data on yield are incom-
plete; nevertheless it is worth while to make a few comparisons based
upon the data available, making due allowance for the evident inac-
curacies in the results.

[Cir. 196]

9

A comparison of yellow pine and blackjack shows an average yield
of 1.8 bushels of cones per tree for the former and 1 bushel for the latter.
This difference is attributable mainly to a difference in the size of
the trees, the yellow pimes on account of their greater age being on
the average much larger than the blackjacks. A comparison of the
classes with respect to the amount of seed produced by a pound of
dry cones gives 1.3 ounces for the yellow pine against 1.7 ounces for
the blackjack. It therefore appears that, although the younger
trees yield less cones, when the amount of seed produced by a given
quantity of cones and the vitality of the seed are considered, these
trees are better seed producers for their size than the older trees.
In comparing the two age groups from 140 to 250 years and from
260 to 400 years we find, as in the case of the blackjack and yellow
pine, a considerably greater yield of cones for the older trees. The
amount of seed produced by a pound of cones is, however, the same
for both-groups. The difference of cone production is much less
pronounced between these two age groups than between the black-
jack and yellow pine, because many of the trees included in the
lower-age class have passed beyond the blackjack stage. Practically
no data are available on trees below 140 years, since the trees were not
felled, and only a small number of cones were picked from each tree.

A comparison of the healthy and unhealthy yellow pine shows only
a slight difference in cone production. There appears a pronounced
difference between the healthy and unhealthy blackjack, but, since
we have accurate data for only two unhealthy blackjacks, this com-
parison is not conclusive. Furthermore, as in the case of germination;
a comparison of healthy and unhealthy trees is of little significance
unless we consider the nature of the injury in the unhealthy trees.

A comparison of the trees classed as unhealthy, considering the
nature of the disease or injury, is shown in Table 3.

TABLE 3.—Comparative yield of cones from healthy and unhealthy trees.

Healthy. Unhealthy.
|

br fay : Ba
¥ y-. |Spike top,| Fire scar, | Heart rot,
Auer ae yellow f yellow : yellow beetle, | 4 yellow

pine. pine. pine.

Yield of cones per tree. bushels. . 1.8 1 0.9 Qu1 2.1 Vet 0.4
Basis oeee tere een cet. trees... Pi 14 6 15 10 3 A

Although the yellow pines affected by heart rot show a yield of
0.3 bushel more than the healthy trees of this class, the difference is
probably due mainly to the fact that the trees affected by heart rot

‘[Cir. 196] |

10

are usually old and consequently of large size. The average diam-
eter of the healthy yellow pine im this study is only 23 inches, against
27 inches for those affected by heart rot. It is probable that, as in
the case of germination, the rot ordinarily has no material effect.

Spike tops show a yield only one-half as great as that gwen for healthy
yellow pines. This is probably due to a reduction in the size of the
living crown. The living branches of spike tops are frequently
heavily laden with cones, and it is probable that the injury causing
the death of the top acts as a stimulus upon cone production, as
upon the vitality of the seeds.

The influence of basal burns upon yield is not fully demonstrated.
The yellow pine bearing fire scars shows an average yield of 0.3
bushel more than the healthy yellow pine; but a comparison of
diameters shows that the scarred trees have an average diameter
of 25 inches against 23 inches for the healthy ones. Whether the
greater yield is due entirely to the greater size of the burned trees
or whether the influence of the burns is also a factor, can not be
determined from the data at hand.

The yield of trees suffering from suppression and attacks of bark
beetle and mistletoe are all much below that of normal trees of ther
class. While the number of trees in each case is too small to furnish
reliable data, the results are very significant.

The yield decreases as the density of the stand increases. ‘The aver-
age yield of cones per tree for open, medium, and dense stands are
1.8, 1.1, and 0.7 bushels, respectively. This difference is undoubtedly
due mainly to a difference in crown development as influenced by
growing space. Since western yellow pine is very light demanding,
the trees growing in close stands have narrow, short crowns, and
only those branches which are exposed to full or nearly full sunlight
bear any cones whatever. Therefore, although the seed produced
by a tree in a close stand is of higher quality than that produced in
the open, as indicated by the germination tests, the total seed-
producing capacity of a tree in a close stand is much lower.

APPLICATION OF CONCLUSIONS.

Blackjacks are the most desirable trees to leave for regeneration
in cutting. The blackjack has the advantage over the older yellow
pine in point of seed quality, and alsoin the rate of growth. A black-
jack left for a future cutting will increase rapidly in volume, while
a yellow pine will increase but slowly or even deteriorate.

In marking for cutting in the yellow-pine type it is the invariable
practice to leave all healthy blackjacks, and also to leave a suffi-
cient number of yellow pines to insure a second cut in about 30 to
50 years. But if a second cut were the only consideration in mark-
ing there would not always be sufficient provision made for seed

[Cir 196] |

11

trees; because the trees left for a second cut do not have to be very
evenly distributed over the area. Thus, one quarter section may
contain an abundance of blackjack and yellow pine suitable to
leave for a second cut, whereas an adjoining quarter section may
contain mostly yellow pine which can be cut now, and very few
seed-bearing blackjacks. Trees below 16 inches in diameter breast-
high usually bear only small quantities of seed, and hence will not
figure materially in the seed crop for the next few years. Therefore,
if immediate reproduction is desired, wherever there are not enough
seed-bearing blackjacks the deficiency must be made up from the
youngest and best developed yellow pines available, even though
not needed for a second cut. The number of trees per acre needed
for seed purposes varies greatly with size and condition; on the
average, from 3 to 5 yellow pines and about twice as many black-
jacks (over 16 inches diameter breast-high) are required.

Spike tops and burned or decayed trees should always be cut,
unless greatly needed for seed purposes. While such trees usually
have a high seed-bearing capacity, to leave them standing until a
second cut can be made involves merchantable deterioration or
total loss. The risk in the case of Jight burns is comparatively
small, but in the case of spike top and decay loss is certain. Since
in extreme cases, however, it becomes necessary to leave defective
trees for want of anything better, it is well to know that, unless too
far gone, such trees can be expected to produce large quantities of
superior seed for a number of years.'

Trees infested by mistletoe or bark beetles, even though not.
lulled, are of little value for seed, and should be cut as being a menace
to the forest. (Instructions in regard to the proper disposal of trees
attacked by bark beetles are given in Bulletin 83, Part I, U.S.
Department of Agriculture, Bureau of Entomology.) An exception
to this rule may be permissible in the case of mistletoe, when the
trees are isolated and when the attack is slight, but not in the case
of bark-beetle infestation. Further investigations in regard to the
effect of mistletoe and bark beetles are needed.

Isolated trees are preferable to those in close stands; first, because
they usually have larger crowns; and second, because they are more
windfirm.

For seed collecting, young trees are most desirable from a purely
technical standpomt on account of the high vitality of their seed;
but in practice, older trees are preferable on account of their heavier
yield, and because the seed from such trees can often be collected
cheaply wm connection with logging operations. Such defects as
spike top, burns, or heart rot are not objectionable; but trees affected
by insects or mistletoe should be avoided.

1 Records have been established to determine how long such trees will persist.
(Cir. 196] O

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