Growth patterns in “knees” of Dawn Redwood in relation to distance from parent tree

Abstract
The dawn redwood (M. glyptostroboides) is one of a few tree species to have roots grow
upright above the ground, commonly referred to as “knees”. Little is known about these knees,
but it is hypothesized that they are used for additional gas exchange for the roots, which are often
submerged in flood waters. This study tested whether the height and diameter of the knees is
related to the distance from a knee’s parent tree. M. glyptostroboides knees were measured at the
Davis Arboretum in Auburn, Alabama. A correlation test was used to determine the relationships
between the height and diameter of the knees compared with the distance from the parent tree. A
correlation was found between the height and distance (p<0.05), but no correlation was found
between the diameter and distance (p>0.1). Additionally, there was a correlation between height
and diameter (p<0.01). The results indicate that the tallest roots are farthest from the parent tree,
suggesting that the outer roots are more likely to participate in above ground gas exchange. This
indicates that M. glyptostroboides that live in areas that are sporadically flooded require more
space to grow than trees that grow on higher ground.

Introduction
Dawn redwoods (Metasequoia glyptostroboides) are one of a few tree species which have
knees: segments of roots which grow in protruding conical structures above ground. The purpose
of the knees remains unknown. Currently, there are no studies related to M. glyptostroboides
knees, however, knees have been studied in the similar species, bald cypress (Taxodium
distichum) (Martin et al., 2015; Kernell et al., 1990; Kramer et al., 1952). The dominant
hypothesis is that the knees assist in providing oxygen to the submerged roots (Martin et al.,
2015; Kramer et al., 1952). Another hypothesis is that they act as stabilizing structures on
account of the trees’ swampy habitats (Martin et al., 2015). There is much observational
evidence that the knees are used in gas exchange, but experimental studies do not display
conclusive evidence (Brown et al., 1984; Martin et al., 2015).
M. glyptostroboides typically grow in low land habitats, such as swamps or otherwise
near bodies of water. (Fan et al., 2020). They are an endangered species native to China, but they
have been successfully introduced to North America and parts of Europe. They are expected to
lose available habitat due to climate change, but habitat loss is mainly driven by changes in
precipitation rather than changes in temperature (Fan et al., 2020). Individual trees may grow
directly near water or on higher ground, but the knees are only grown on individuals grown in
areas with sporadic flooding and exposure of soil; therefore, knees do not grow on trees in deep
water or on trees completely out of water (Kramer et al., 1952; Kernell et al., 1990). The height
of the knees is related to water depth when the area is flooded: the tallest knees are found in the
deepest flood waters (Kernell et al., 1990). This supports the theory that the knees are searching
for areas with higher oxygen to provide that oxygen to the submerged roots. The wood of the
knee is porous and the tips lack thick bark, which could prevent the inhibition of gas diffusion
through the knees (Martin et al., 2015).
The purpose of this study is to build upon the little evidence known about the
characteristic of trees growing knees, especially for the species M. glyptostroboides. Contrary to
prior research, the height of M. glyptostroboides knees growing in the Davis Arboretum on
Auburn University’s campus do not appear to be correlated with water depth. Instead, the height
appears to be related to the distance from the parent tree. I hypothesize that the larger knees are
farthest from the parent trees because the outer roots are searching for adequate areas above
ground to assist in gas exchange. I predicted that as the distance from the parent tree increases,
the height and diameter of the knees will significantly increase.

Methods
The location of the M. glyptostroboides used in this study was the Davis Arboretum on
Auburn University’s campus in Auburn, Alabama. The trees grow in damp soil near a small
pond and stream. There are few other plants around the M. glyptostroboides, indicating that the
low ground level does flood.
The knees of four M. glyptostroboides trees were measured in the fall of 2020. Since the
knees are slightly curved towards the tree that they grow from, I was able to determine the parent
tree of each knee (Kummer et al., 1991). The height of the knees was measured from the base of
the knee at soil level to the topmost tip of the knee. The diameter was measured approximately at
half-height of each knee in order to approximate the average diameter. Distance was measured
from the parent tree to the middle of the tip of the knee.
Measurements were taken in inches, and converted to centimeters. A correlation test was
used to analyze the data with correlation graphs for height related to distance and diameter
related to distance. An additional correlation test was used to analyze diameter related to height.

Results
A correlation was found between the height and distance from the parent tree, but not for
diameter and distance from parent tree. The height of the knees in relation to the distance from
the parent tree resulted in a correlation coefficient of 0.346 (p<0.05), with an average of 27.98cm
and standard deviation of 18.07 (Figure 1). The diameter of the knees in relation to the distance
from the parent tree resulted in a correlation coefficient of 0.152 (p>0.1), with an average of
19.63cm and a standard variation of 7.36 (Figure 2).

90
80
70
60
Height (cm)

50
40
30
20
10
0
0 50 100 150 200 250
Distance from Parent Tree (cm)

Figure 1. The relationship between the height of the M. glyptostroboides knees and distance from
its parent tree collected at the Davis Arboretum in Auburn, AL in September 2020.
 40
35
30
Diameter (cm)

25
20
15
10
5
0
0 50 100 150 200 250
Distance from Parent Tree (cm)

Figure 2. The relationship between the diameter of the M. glyptostroboides knees and the
distance from its parent tree collected at the Davis Arboretum in Auburn, AL in September 2020.

The correlation test for the diameter and the height of the knees resulted in a correlation
coefficient of 0.542 and a p-value less than 0.01.

Discussion
Because statistical analysis of height related to distance resulted in a p-value less than
0.05, it can be concluded that the height of the M. glyptostroboides knees is correlated with the
distance from its parent tree. However, since the statistical analysis for the diameter of the knees
resulted in a p-value greater than 0.1, it can be concluded that the diameter of the knees is not
related to the distance from the parent tree. The statistical analysis of diameter related to height
resulted in a p-value less than 0.01, so it can be concluded that the diameter and height are
directly correlated. This means that the tallest knees are the ones farthest from the parent tree, but
a specific diameter does not relate to a certain distance.
These results contradict the results of past studies, which show that the height of the
knees was related to the depth of occasional floodwaters (Kernell et al., 1990). The correlation
between the height of the knees and the distance from the knee’s parent tree could demonstrate
that it is the outer roots that are more involved in gas exchange (Kramer et al., 1952, Martin et
al., 2015). As the roots grow outwards from the tree, the roots farthest from the tree potentially
exhibit a lack of oxygen, and grow upwards in search of oxygen. However, there often are knees
that grow close to the parent tree; these knees likely require less oxygen from above the surface
than the knees farther from the parent tree. Another explanation could be that the outer roots
require additional oxygen sooner than the closer roots in relation to the age of the tree. If this is
the case, the height of the knees would be directly related to the age of the knees implying that
the smaller knees are smaller because they have not been growing for as long as the taller knees.
Provided that the tallest knees grow the farthest from the parent tree, this could imply that the M.
glyptostroboides that require additional gas exchange would also require additional space to
grow. This could be important knowledge because M. glyptostroboides is an endangered species,
and the greatest threat to the species is habitat loss.
The knees measured at the Davis Arboretum grew on uneven ground, which could affect
the resulting measurements of their heights, possibly making some knees appear shorter than
others. This would explain the apparent randomness of the data points in Figure 1 despite the
statistical analysis showing correlation. Also, the diameter was measured at the knees’ half-
height, when the area of greatest diameter could have been measured instead. An additional
study might measure diameter at the widest point of the knee in relation to the distance from the
parent tree.
Future studies should take into account the unevenness of the ground if necessary, as well
as the age of the knees. This would provide more accurate data in a replication of this study.
Additionally, measuring the amount of gas exchange related to the height of the knees would
provide further knowledge on their use and growth patterns.

References
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domes in north-central Florida."

Fan, Kexin, et al. (2020) "Do climate and human disturbance determine the sizes of endangered
Metasequoia glyptostroboides trees in their native range?" Global Ecology and Conservation 21:
e00850.

Kernell, Judith Leeds, and Gerald F. Levy. (1990) "The relationship of bald cypress knee height
to water depth." Castanea: 217-222.

Kramer, Paul J., Riley, Walter S., and Bannister, Thomas T. (1952) "Gas exchange of cypress
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Kummer, Hans, et al. (1991) “Nutritional exploitation of dead trunks: another function of cypress
knees (Taxodium distichum)?” Trees 5:122-123.

Martin, Craig E. and Francke, Sarah K. (2015) "Root aeration function of Baldcypress knees
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