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Soybean and Bees

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sible that the nectar production dynamics of a species can be determined. The strategy of<br />

the plant of offering nectar, the activity patterns, frequency <strong>and</strong> diversity of pollinators of a<br />

plant species, the rates of nectar consumption by animals, among others, could not be understood<br />

without deep knowledge of nectar characteristics <strong>and</strong> dynamics.<br />

Nectar production <strong>and</strong> the role of enzymes<br />

Several studies targeted the action of the apoplastic invertase on tissues surrounding the<br />

phloem, which create the flowing of sugars to non-photosynthetic tissues, as is the case of<br />

flowers. The apoplast is the free diffusional space outside the plasma membrane, interrupted<br />

by air spaces between plant cells <strong>and</strong> by the plant cuticle. This way, the apoplast is formed<br />

by the continuum of cell walls of adjacent cells as well as the extracellular spaces, forming<br />

a tissue level compartment. The apoplastic route facilitates the transport of water <strong>and</strong> solutes<br />

across a tissue or organ. This process is known as apoplastic transport. The apoplast is<br />

important for all the plant’s interaction with its environment. The main carbon source (carbon<br />

dioxide) needs to be solubilized in the apoplast before it diffuses through the plasma<br />

membrane into the cytoplasm <strong>and</strong> is used by the chloroplasts during photosynthesis. The<br />

apoplast is also a site for cell-to-cell communication.<br />

Ruhlman et al. (2010) observed that the production of nectar is closely linked to the presence<br />

of the enzyme apoplastic invertase, therefore, larger nectar production was dependent on<br />

an increase in enzyme activity in addition to the photosynthetic capacity of the plant itself.<br />

Then we can infer that increasing the volume of nectar promoted by its withdrawal is largely<br />

linked to an increased activity of the invertase enzyme. These same authors observed that<br />

in modified Arabdopsis, silencing the gene encoding one of the isoforms of the predominant<br />

enzyme in the reproductive organs prevented the production of nectar.<br />

The rationale of this process was proposed by Cheng <strong>and</strong> Chourey (1999). Invertase promotes<br />

the hydrolysis of sucrose to the hexoses glucose <strong>and</strong> fructose <strong>and</strong> can be located in the cell<br />

wall (apoplastic), vacuole (vacuolar) or cytoplasm (cytoplasmic). The cell wall invertase is<br />

important when unloading the phloem apoplastic follows a path facilitating the passage of<br />

sucrose to the drains tissues. The authors’ studies concluded that the importance of apoplastic<br />

invertase is not restricted to the breakdown of the sucrose molecule, since the injection<br />

of glucose <strong>and</strong> fructose has the same effect on the passage of photosynthates to the phloem<br />

tissue drains. In this case, the authors attribute to the enzyme a possible broader role as signaling<br />

or regulatory factor.<br />

Further evidence of the importance of apoplastic invertase in phloem unloading process<br />

that follows this path is the abundance of transcripts in these tissues, according to Jin et al.<br />

64 SoybeAn <strong>and</strong> bees

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