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SOURCE/SINK RELATIONSHIPS DURING TUBER GROWTH 1

Robert B. Dwelle 2
Carbon partitioning and sink/source interactions in plants have been
the subject of numerous reviews (e.g., 1, 19).

Definition of 7~rms
"Source" refers to plan.t tissues which are net carbon producers, generally green, photosynthetic tissues. Source tissues in a potato plant are
primarily leaves and stems. A "sink" is a net carbon user, which includes
respiration, growth and storage ofcarbon compounds. Sink organs include
tubers, roots, rapidly growing tissues, or even old and shaded leaves. In
a potato plant, the sink of greatest interest is the tuber. "Sink strength" or
"sink demand" refers to the ability of sink organs to attract or accumulate
carbon compounds.

Carbon Flow
The movement of carbon compounds through the plant can be referred to as "carbon flow; and for convenience we can divide the process
of carbon flow into 5 components. The first is carbon fixation or photosynthesis.
The second phase is partitioning of newly-fixed carbon within the leaf
between sugar and starch. If carbon is partitioned to sucrose, it is available for rapid export from the leaf tissue. Starch, on the other hand, is accumulated within the leaf, but it can be remobilized during the night.
Moorby (12) found that within the starch grains of the potato leaves, starch
is laid down in concentric shells and later removed in the reverse order.
In other words, starch producedjust prior to a dark period will be the first
mobilized during the subsequent dark period. This pattern of starch deposition and removal can obscure short-terrn patterns of photoassimilate partitioning. Nevertheless, Moorby (11) was able to demonstrate that foUowing
tuberization, source leaves export greater quantities of current photoassimilates than before tuberization. Oparka et al. (14), have identified a longerterna storage pool of starch in the leaves that may be available to both tubers
and leaves over a period of several weeks.
The third phase of carbon flow is phloem loading of sucrose, which
is an active process (requires energy). The exact nautre of the loading proc
is not yet known, but is being investigated currently in several laboratories.
'Approved for publication by the Director of the Idaho Agricultural Experiment Station
as research paper No. 90751.
2Professor and Chairman, Plant Science Division, Unir. of Idaho, Moscow, ID 83843.
Accepted for publication September 19, 1990.

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The fourth phase is long-distance translocation, which is thought to

be driven by mass flow. Transport ofcarbon compounds is rarely limiting;
most plants have more phloem than needed. Moorby (12) has maintained
that the capacity of the phloem sieve tube cells in potato plants is generally n o t a limiting factor in photoassimilate tra.nslocation. Rather, the supply
of materials from the source leaves and/or the removal of these materials
at the sinks are the critical control points.
The fifth phase is phloem unloading and uptake/utilization by the sink
tissues. An active sink must maintain low concentrations of sucrose at the
site of phloem unIoading. This is reportedly an active, carrier-mediated
process (15), and cell turgor appears to influence both sucrose uptake and
conversion of sucrose to starch (16). It has been suggested that sucrose synthase plays an important role in determining the relative sink strength of
potato tubres (17, 20). The subject of tuber growth and metabolism will
be covered in the next paper of this symposium.
To summarize the carbon flow process, the classical view holds that
a sucrose gradient between source leaf and sink determines direction and
extent of movement ofsucrose. Sinks purportedly influence patterns ofcarbon distribution by regulating unloading, thus increasing the sugar concentration gradient between source and sink (7).
Source-Sink Interactions

Numerous reports with varied crops have shown that increased sink
demand can result in increased source output, decreased sink demand can
result in decreased source output, and partial removal of the source Ieaves
can result in increased output by the remaining leaves [see review by Dwelle
(5)]. Similar source-sink interactions have been shown with potatoes. Nosberger and Humphries (13) reported that removing potato tubers reduced
the plants' net photosynthesis rate. Moorby (11) found that after tuber initiation, a two- to three-fold increase in photosynthetic rate occurs and the
proportion of assimilates exported from the leaf is doubled, most of this
going to the tubers. Moll (10) reported minimum photosynthesis rates at
the time of tuber initiation, a maximum during the period of linear bulking of the tubers, and then declining rates at the time when the crop reached
60-70% of the maximal yield. Dwelle et al. (4), found a similar pattern in
fietd studies.
Cranshaw and Radcliffe (3) found that defoliation of 67% of the potato
canopy early in the season resulted in only slight yield reduction, suggesting that an increased output by the remaining source tissues may have compensated for the lost leaves. Ewing et al. (6), have reported similar results,
including compensatory growth and increased rates of photosynthesis in
response to partial defoliation.
Some reports in the literature have not found source-sink interactions
with certain potato cultivars and under certain conditions (2, 5). In addition, we have found under greenhouse conditions that when all tubers are

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METABOLISMSYMPOSIUMPAPERS

831

removed from the potato plants, the mberization stimulus can be so powerful
that numerous axillary tubers will form on the above-ground stems. Under these circumstances, alternate sinks were formed and source activity
(photosynthetic rate) was undiminished (Dwelle, unpublished). Nevertheless, it has been shown that source-sink interactions can occur with many
potato cultivars and under many conditions.

Coordination of PhotosynthesisAnd Sucrose Synthesis in Leaves

As an illustration of the complexity of control mechanisms, several
laboratories are investigating interactions between photosynthesis and sucrose synthesis in leaf tissues. Fructose-2,6-bisphosphate has been identified a s a signal metabolite (rather than ah intermediate) that senses how
much photosynthate is available to make sucrose and adjusts the rate of
sucrose synthesis accordingly (18). For example, rapid photosynthesis lowers
the level of fructose-2,6-bisphosphate, which activates sucrose-P synthase
(SPSase), thus promoting sucrose synthesis. To further illustrate the complexity, activity of SPSase varies depending on genetic components, photoperiod, nitrogen source, developmental stage of the plant, carbon dioxide
concentration, water stress, and so on. Synthesis and degradation of fructose2,5-bisphosphate is regulated by metabolites such as DHAP, PGA, Fru-6P, and inorganic phosphate (8). Obviously, the control mechanism has not
yet been worked out, but the pieces are slowly being assembled.

Genotypic Variationsin Sink/Source Interactions

Li and Kao (9) have reported interesting genotypic variations with
respect to source potentials and sink capacities of sweet potatoes. In this
instance the primary sink is a tuberous root rather than a true tuber, but
the types ofvariations they have found may be similar in SolanumtuberosumL.
Li and Kao tested 20 genotypes and found significant differences in
source potential and in sink capacities among the tested genotypes. They
performed a number of reciprocal grafts, grafting the top ofone genotype
to the root of another. A s a result of these reciprocal grafts, they were able
to characterize the source potential and sink capacity of each genotype, and
they found every combination possible. Some of the high-yielding genotypes had both high source potential and high sink capacity. Two genotypes
had high source potential but poor sink capacity. Some of the low-yielding
genotypes were poor in both source potential and sink capacity. They also
found that some genotypes with high source potential often did not respond
to variations in sink strength; in other words, some genotypes with high
photosynthetic rates continued these high rates no matter how strong or
weak the sink activity. This information raises exciting possibilities for the
bio-engineering of new cultivars.
For years we have debated in the literature whether or not potatoes
(Solanum tuberosum)exhibit true source/sink interactions. Perhaps the answer
is, "Sometimes yes, and sometimes no: The work of Li & Kao was done

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with sweet potatoes; it would be interesting to conduct the s a m e experim e n t s with Solanum tuberosum.

Concluding Questions
T h i s review has u n d o u b t e d l y raised m o r e questions t h a n it has answered. Two questions in particular need attention. T h e first: " W h a t is the
control m e c h a n i s m ? " H o w does sink influence source (or vice versa)? A r e
h o r m o n a l factors involved? O r is it a direct feedback m e c h a n i s m ? A seco n d m a j o r question is: "What are the sink/source interactions in potatoes?"
T h e y need to be characterized for m a n y genotypes. We need to d e t e r m i n e
if the system is sink-driven, source-driven, or some of each. T h e u s e of
reciprocal grafts could p r o d u c e interesting i n f o r m a t i o n to help c h a r a c t e r ize genotypic variations.
Literature Cited

1. Bangerth, F. 1989. Dominance among fmits/sinks and the search fora correlative
signa]. Physiol Plant 76:608-6t4.
2. Burton, W.G. 1989.The Potato, ThirdEdition. John Wiley & Sons, Inc., New York, NY.
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3. Cranshaw, W.S. and E.B. Radcliffe. I980. Effect ofdefoliation on yield ofpotatoes.
Jour Econ Entomol 73:131.
4. Dwelle, R.B., G.E. KleinkopfandJJ. Pavek. 1981. Sotmatal conductance and gross
photosynthesis of potato (Solanum tuberosum L.) as influenced by irradiance, temperature and growth stage. Potato Res 24:49-59.
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1965. The influence of removing tubers on drymatter production and net assimilation rate ofpotato plants. Ann Bot 29:579-588.

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14. Oparka, K.J., B. Marshall and D.K.L. Mackerron. 1986. Carbon partitioning in a
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C onference of the European Association of Potato
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