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Soybean and Bees

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crose, fructose, <strong>and</strong> glucose; dominant fructose <strong>and</strong> glucose. It has been proposed that nectar<br />

quantity <strong>and</strong> quality may play an important role in determining the search of foraging honeybees<br />

(Waddington <strong>and</strong> Holden, 1979), although it is not clear whether bees alter their<br />

foraging strategies considering the hourly <strong>and</strong> daily fluctuations in nectar characteristics of<br />

individual species.<br />

Frey-Wyssling (1955) supported the theory that nectar secretion is related to the phloem<br />

supply to floral nectaries because the number of cells between phloem sieve tubes <strong>and</strong> the<br />

surface of floral secretory tissue cannot exceed a certain amount (in most cases about ten<br />

cells) for abundant nectar secretion to occur. However, nectar composition differs considerably<br />

from that of phloem sap (Ziegler et al., 1959) <strong>and</strong> has been proposed by Luttge (1977)<br />

to be a function of specific active transport mechanisms.<br />

In this context, the primary enzyme would be the apoplastic invertase located in the tissues<br />

surrounding the phloem unloading <strong>and</strong> that allows sucrose to non-photosynthetic organs<br />

that follow an apoplastic pathway after the phloem, such as the reproductive organs. Ruhlman<br />

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

the enzyme apoplastic invertase; therefore, a greater nectar production was dependent on<br />

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

These authors observed that in Arabdopsis, in the absence of genes encoding isoforms of the<br />

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

Invertase promotes the hydrolysis of sucrose to the hexoses glucose <strong>and</strong> fructose <strong>and</strong> can be<br />

located in the cell wall (apoplastic), vacuole (vacuolar) or cytoplasm (cytoplasmic). The cell<br />

wall invertase is important when unloading the phloem follows an apoplastic path facilitating<br />

the passage of sucrose to the drains tissues. Some studies even conclude that the importance<br />

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

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

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

as signaling or regulatory (Cheng <strong>and</strong> Chourey, 1999).<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 (Jin et al., 2009) <strong>and</strong> its<br />

almost non-existence in tissues with symplastic unloading (Ruan <strong>and</strong> Patrick, 1995), both<br />

observed from tomato tissues. In Vicia faba, tissues surrounding the seed formation, which<br />

have no connections of plasmodesmata, also show high expression of the enzyme (Weber<br />

et al., 1996).<br />

Honeybees collect sucrose solutions in the 30-50% range in preference to higher or lower<br />

concentrations (Waller, 1972; Woodrow, 1968). Jamieson <strong>and</strong> Austin (1956) found that<br />

SoybeAn <strong>and</strong> bees<br />

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